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Full text of "The genus Clusia section Criuva (Clusiaceae) in Guyana"

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\jP Biodiversity 

SIDA, contributions to botany. 

[Dallas :W. F. Mahler, etc.]1962- 1 3 

V.16 (1994): 
Page(s): Page [649], Page 650, Page 651 , Page 652, Page 653, Page 654, Page 655, Page 
656, Page 657, Page 658, Page 659, Page 660, Page 661 , Page 662, Page 663, Page 664, 
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673, Page 674, Page 675, Page 676, Page 677, Page 678 

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Botanical Research Institute of Texas 

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Fort Worth, TX 76102-406(1 USA. 


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San Gregorio, CA 94074, US. A, 


The genus Clmia section Cr'mva is revised for species within the G nay ana Florist ic Pnn- 
ince. Eleven species are recognized, of which five, Cinua grammademmdes Pi poly, C. 
multdtnmta Pipoly, C asymmetriai Pipoly, C magutredna Pipoly, and C. guayanae Pipoly are 
described as new, illustrated, and their phylogenetic relationships are discussed. Amplified 
descriptions are provided for the section and each hitherto known species, along with keys 
to the species within the section, and citation of representative specimens. Clusia reducia is 
reduced to synonymy under Clusia opaca and Clmia h^xmarpa subsp. ptarmpumna is re- 
duced to synonymy under C hi^xaairpu. 


Se presenta una revision caxonomica del genero Clusia seccion Cnuva para las especies 
procedentes de la Provincia floristica Guayanesa. Se reconocen once especies, dentro de las 
cuaies hay cinco, Cluua gvamnmdmifjides Pipoly, C multiiuimtd Pipoly, C. asymmetyka Pipoly, 
C. magiureand Pipoly, y C gi^ayanae Pipoly que se describen como nuevas, se i lust ran y se 
discutc su parentesco. Se ofrecen descripciones ampliadas para la seccion y especies 
previamente conocidas; tambien se presentan claves para identificar his especies dentro de 
la seccion, y un listado de especimenes representativos, Se reduce Clusia redmta a hi sinonfniia 
bajo C. opacd y C. hexacarpa subsp. ptaniepKuziui bajo C. hexacarpd, 


While preparing a diagnosric treatment of the genus Clusia L. for che 
Flora of the Venezuelan Guayana and a more comprehensive one for Flora de 
Colombia^ section Crtuva was revised. This synopsis treats the species of 
section Criuva indigenous to the Guayana Crystalline Shield (sensu Magu- 
ire 1979), and provides descriptions, synonymy, specimen citations distri- 
butions, hypotheses of phylogenetic relationships, ecology and conserva- 
tion status for each Guayana species of the section. 

Section Crtuva is closely related to sections StaimKlusia Planch. & Triana 

SiDA 16(4): 649-678. 1995 


Si PA 16(4) 

and Clmiastrum Planch. & Triana, (Engler 1888, 1893; Vesque 1893), but 
easily distingLiished from both by the stami nodes with well-developed an- 
thers, abruptly widened and connivent basally. While Engler (1888, 1893) 
felt that Criuva wa5 most closely related to Crmvopsis Planch. & Tnana, his 
classification was based mostly on petal number rather than on some quali- 
tative feature, Pipoly and Graff (1995) have indicated that the most closely 
related group to section Criuvopsis is section Bf'achystemon Engler. Because 
the formats of the floristic treatments in preparation do not allow for com- 
plete descriptions and synonymy, the present treatment is intended to pro- 
vide data for the Guayana taxa over their entire range, until a more com- 
prehensive revision is prepared, 


Clusia L. section Criuva Planch, & Triana, Ann. Sci. Nat. ser 4, l4: 325. 

Clnsia still. Crnwa subsecr. lliicrtHva Engler in Marrius, Fl. Bras. l2(i):106. 1888. 
Clusid sLib^i^cnus Criuva Btrirh. & Hook, section Encrunkt (Planch. & Triana) Engler, 
Pflanzenfam,3(6):225. 1895. 

Free-standing terrestrial shrubs, trees or hemiepiphytes; hirex cream ish 
yellow to translucent, when translucent, oxidizing to yellow. Leaves sessile 
or petiolate; petioles canaliculate, marginate or trigonal. Inflorescence a 
terminal pyramidal panicle of cymes, a reduced compound axillary cyme or 
rarely, a solitary terminal flower, often subtended by fbliaceous bracts; sec- 
ondary inflorescence bracts car nose or cartilaginous; bracteoles 2-4(-l4), 
coriaceous, carnosc or cartilaginous. Staminate flowers with spreading pe- 
rianth; sepals 4-8, coriaceous or chartaceous, the outer opposite, the inner 
decussate, imbricate or contorted; petals 4—6, coriaceous, carnose or carti- 
laginous, rhe outer opposite, the inner decussate, imbricate, or contorted, 
usually progressively smaller acropetally; androphore recranguloid, 
pentagonoid or rarely cubic; stamens numerous, the filaments apically free, 
connate at the base, fleshy, often thickened at the base; the filaments flat, 
anthers muticous, rounded or emarglnate, equalling or shorter than the 
filaments, the thecae clavate, oblong or linear, slightly longer than the con- 
nective, dehiscent by longitudinal slits; pistil lode absent or rarely, vesti- 
gial, usually without resin. Pistillate flowers as in stammate; stami nodi a 4- 
numerous, free, linear, with vestigial anthers; ovary 4—8 locular, one to 
multiovulate, ovule placement variable; styles obsolete or equalling loc- 
ules in number, free, terete or angled, thickened, radiate, stigmas subpeltate, 
orbicular, ovate, cuneiform or pentagonal Fruit a dehiscent capsule, ovoid, 
oblongoid, prismatic or fusiform, stigma and styles persistent. 

TypI: Sphcihs: CJhsui o'hna Cambessedes in A. Sr.-Hilaire, Fl, Bras, Merid, 1:245, 1825. 

PrpOLY AND Graff, Clusia section Criuva in Guayana 651 


1. Leaf blades chartaceous to sLibcoriaceous, dull above, pallid below, the api- 
ces acute to acuminate, bearing round, magenra glands abaxially. 
2. Apical portions of branchlets decussately alate; lea\^es sessile or on broadly 
marginate; petioles, 0.5-1.0 cm long. 

3. Leaves 1.5^,0 cm wide, sessile; peduncle 2.0-3-5 cm long; sepal mar- 
gins opaque; entire 1* C. clid^me 

3- Leaves 2—6 cm wide, on broadly marginate petioles, 0.5-1 cm long; 
peduncle obsolete to 3 mm long; sepal margins scarious, hyaline, in- 

cised-erose 2. C, gra^muadmiokks 

2. Apical portions of branchlets subterete to tetragonal, the corners rounded, 

not alate; leaves on narrowly marginate or trigonal; petioles 1-3.5 cm 


4- Branchlets subterete or weakly tetragonal; leaf blades symmetric, li- 
ear latex canals conspicuous; petioles marginate, thin, 1-1.5 cm long; 
sepals 7, without conspicuous latex canals, the outer sepals acute to 
acuminate apically; petals decussate, thickly carnuse, suborbicular; an- 
thers obovoid,,... ^...^ " 3. C. muhdimata 

4. Branchlets strongly tetragonal; leaf blades asymmetric, linear larex ca- 
nals inconspicous; petioles trigonal, thick, 2-3.5 cm long; sepals 4, 
linear latex canals conspicuous, tlie outer sepals rounded apically; pet- 
als imbricate, coriaceous, obovate; anthers ovoid .4, C. asymmetrka 

1. Leaf blades stiffly coriaceous to cartilaginous, nitid at least above, the apices 
obtuse, broadly rounded or truncate, without round magenta glands abaxially. 
5. Leaf blades coriaceous, rarely thickly so, but the secondary veins promi- 
nent above and below; inflorescence erect, the rachis flattened; bracteoks 
numerous, small, similar to the sepals; flowers numerous. 
6. Petioles broadly marginate, 3-10 mm long; leaf blades obovate to very 
widely obovate, the bases cuneate, fully decurrent to petiole base; in= 
florescence rachis longitudinally ribbed, not alate; stigmas on persis- 
tent styles, ca. 1 mm long; fmit ovoid at first, then fusiform at maturity 
, „ ,..,,,..... ,..,,..,......,,..,...... 5 . C mekbiorii 

6. Petioles canaliculate, 15-25(-28) mm long; leaf blades oblong, ellip- 
tic or rhombic, the bases obtuse, not decurrent on petiole; inflorescence 
rachis narrowly alate; stigmas sessile; fruit subglobose at first, then 

globose at maturity ,,. ...^ 6. C. vardomre 

5. Leaf blades thickly coriaceous to cartikiginous, the secondary veins not 
easily discernible without magnification; inflorescence pendent, the rachis 
not flattened; bracteoles few, smaller than the sepals; flowers solitary or 
in a few-flowered, pyramidal compound cyme. 

7. Petioles broadly marginate; flowers solitary (-3) or in reduced, 6- 
flowered cymes; stigmas on persistent styles; fruits ovoid to subglobose, 
strongly ribbed. 

8, Petioles, 6-15 mm long; flowers terminal, solitary; petals obovate 

to orbicular; staminodes 8-9^ 

9.Leaf blades 2.5^.5 cm wide; bracteoles 3.5-4 mm long; sepal 

margins not scarious; petals 6; carpels 6—8... 7. C. hexacarpa 

9. Leaf blades 5-7 cm wide; bracteoles 12-14 mm long; sepal mar- 
gins scarious; petals S; carpels 7—10 8. C radiata 

652 Sjda \6(4) 

8. Petiole obsolete co 6 mm long; flowers in an axillary, branched, 3- 

6-Howcred cyme; petals suborbicular; scan^iiiiodes 4 ,...../.... 9^ C. maf^///ream/ 

7. Petioles canaliculate; flowers in 6-18-flc)wcred cymes; stigmas sessile 

to subsessile; fruits globose, smootb, without ribs, 

10. Leaf blades obtuse to broaLily roundi^d basally; peduncle 3-5 mm 
long; sepals 4^-6, suborbicular; petals coriaceous; stanii nodes nu- 
merous; stigmas orbicular, sessile. 10, C\ opaa^ 

10. Leaf blades broadly acute to obtuse basally; peduncle 5-12 mm 
long; sepals 2, oblate; petals cartilaginous; staminodes 4; stigmas 
pentagonal, subsessile ,. 1 1 . CV. guayamre 

1. Clusia duidae Gleason, Bull Torrey Bot. Club 58:402. 1931. Typp. VF.N- 

EZUELA. Ama/onas: Cerro Duida, slopes of ndge 25, 1,800-2,200 m, (fr), Td!e 
429 (HOioTYPf:: NY!; lsotypl: F!). 

Shrub or small tree to 4 m tall; branchlets decussately short-akitc, the 
wings 1-1.8 mm wide. Leaves petiolate; blades charraceous to subcoria- 
ceous, oblong or oblanceolate, 6-8 cm long, {L4-)2.1-4 cm wide, apex 
acute, base abruptly acute, symmetric, dull above, pallid and bearing round, 
magenta glands below, midrib slightly prominent above, prominently raised 
below, secondary veins numerous, prominently raised above and below, 
united by a submarginal collecting vein, the margin flat, entire, glabrous.; 
petioles obsolete. Inflorescence temiinal, erect, pyramidally paniculate, 5- 
7(-8) cm long, 6-9 cm wide, the flowers cymose; peduncle 2.0-3O cm 
long; secondary bracts coriaceous, ovate, 2-5 mm long, apex acute, cari- 
nate; bracteoles 2, coriaceous, cordate, 2.4-2.6 mm long, 2.5-2.7 mm wide, 
apex acute to obtuse, base somewhat auriculate, carinate, the margin en- 
tire, opaque; not scarious; pedicels angulate, 6-12 mm long. Staminare 
flowers unkown. Pistillate flowers yellow; sepals 6, coriaceous, suborbicu- 
lar, 4-6 mm long, 5^2-6 mm wide, apex broadly rounded, carinate, linear 
latex canals few, inconspicuous, the margin entire, hyaline, scarious; petals 
5, coriaceous, acropetally smaller, obovate, the outer pair opposite, 4^.5 
mm long, 3-3.5 mm wide, apex broadly rounded, somewhat cucultare, 
linear latex canals lew, inconspicuous, the margin entire, opaque, not scari- 
ous; staminodes 4, linear, 2.2-2.5 mm long, the filaments 1.2-1.5 mm 
long, L3 mm wide at base, the anthers linear, 1 mm long, apex muticous, 
base obtuse; pistil oblong, 3-3.5 mm long, 2-2.5 mm diam., carpels 4; 
styles angular, thick, ca. 3 mm long, stigmas orbicular, L6 mm long and 
wide. Fruit prismatic, 15-30 mm long, 6-8 mm wide, costate. 

D/j7r/i////o;L— Endemic to the Guayana Crystaline Shield, Duida- 
Marahuaca Subprovince of Maguire (1979), at 1,225-2,200 m elevation. 

Ecology and consaTci lion status. — Ciusia duidae occurs in mossy elrtn forest 
formations, where it forms mats of interwoven stems. Libel data indicate it 
IS locally common, and the fact that its known ramre fills within national 
parks indicates that it is probably not threatened. 

PiPOLY AND Graff, Clusia section Criiiva in Guayana 653 

Representative specimens examined: VENEZUELA, Depto. Atabapo: slope of Cerro 
Marahuaca, upper Rio Yameduaka, 03"38^N, 65"28'W, 1,225 m, 17-18 Feb 1985 (fr), R. 
Lksner 17603 (BRIT, MO, VEN); Cerro Marahuaca, SE slopes, below Salto Los Monos on 
tributary of Rio Iguapo, at headwaters, 03^35'N. 65^23'W, 1,500-1,600 m, 1) Mar 1985 
(pisr. fl, fr), II Liesner 18533 (MO, VEN), 20 Oct 1988 (fr), II Lksner 25133 (BRIT, MO, 
VEN); Cerro Marahuaca, SW-facing slopes, headwaters of Ri'o Iguapo, S sector of the SE 
mounrain, 03^36^00" N, 65^23^10" W, 1,560 m, 13-14 Oct 1983 (pist. fl, fr)J. Sleym/u^rk 
129648 (BRIT, MO, VEN); Cerro Huachamacari, Rio Cunucunuma, vicintiy of summit 
camp, 1,800 m (pist. fl), B, Maguire et al 3()()I7 (F, NY, US, VEN). 

Clusia duidae appears to be most closely related to C. grammadenioides ^ 
but is easily cliscinguished by its sessile leaves with abruptly acute leaf 
bases, entire sepals, and smaller inflorescence, 

2. Clusia graminadenioides Pi poly, sp. nov. (Fig. I) 

Ob ramulurum internodiis quadrangiilatoscjue alatos, folia scssila atque canalis resmiferis 
permanifeste praeditis C. Jmckie vaide affinis, sed ab ea ramulis later (non anguste) alatos, 
folio rum bases sensim largo decrescenres (nee abrupte acutes), petiolos late marginatos 5— 
10 mm longos (non obsoletos), sepal is secus marginem eroso-dentatis (nee integerrimis), 
pro mp tern dignoscenda. 

Hemiepiphyte, attaching to tree trunks by adventitious roots; branchiets 
tetragonal, broadly and decussately alate, 3-5 mm diam^, the wings 2-8 
mm wide, (therefore the stem appearing 8-13 cm diamj, the wings thinly 
cartilaginous, glabrous. Leaves petiolate; blades subcoriaceous, narrowly 
elliptic to obovate, (9-5-)l4-l 9(-25) cm long, (2-)2,5-5(-6) cm wide, 
apically broadly to narrowly acute, basal ly long-attenutate, decurrent on 
the petiole, symmetric, bearing round magenta secretory glands below, 
without conspicuous latex canals, midrib raised above and below, second- 
ary veins numerous, prominent above and below, united by a prominent 
submarginal collecting vein, glabrous, the margin revolute, thin, entire; 
petioles broadly marginate, 0.5-1 cm long, glabrous. Staminate inflores- 
cence erect, terminal, tripinnately pyramidal paniculate, 8—9 cm long, 11— 
17 cm wide, the flowers cymose; peduncle obsolete to 3 mm long; bracteoles 
2, coriaceous, oblate, K8-2 mm long, 2.3-2.5 mm wide, apex rounded to 
obtuse, margin scarious, the band to 0.3 mm, incised with 2-3 incisions, 
glabrous; pedicel tetragonal, 2,2 mm long, glabrous. Staminate flowers 
white, cymose; sepals 8, chartaceous, dimorphic, the outer whorl 2, oppo- 
site to the bracteoles, oblate, 2.5-2.6 mm long, 3-3.2 mm wide, apex 
rounded, somewhat cucuUate, latex canals consipuous, linear, submarginal, 
the margin scarious and hyaline, variously incised-erose, appearing serrulate- 
dentate, the border of 0.5 mm wide, glabrous, the inner whorl 6, imbri- 
cate, chartaceous, suborbicular to oblate 3^8-4.2 mm long, ^5-6 mm wide, 
apex obtuse, cucuUate, the latex canals lineate, prominent, submarginal, 
the margin hyaline, erose-fimbriate; petals 5, contorted, coriaceous, acro- 
petally smaller, oblong to widely ovate, 4.2^.6 mm long, 2,4-3.4 mm 


SiDA 16(4) 

F[c;. 1 C///sia grammukr/huh Pipoly, A, Habit, sliowing the aktc bnuKhlets, B. 2-ll{nvered 
cyme. C Sepals, showint^ the sciirious, hyahne, incised -erase margins. D. Petals, showing 
the linear larex canals, entire margins, and cuculkue habit. E. Androphore, and adroecium, 
A-E, tirawn from type. 

PrpoLY AND Graff, Ciusia section Criuva in Guayana 655 

wide, apex rounded, highly cucuUate, not carinate, latex canals very promi- 
nent, marginal, the margin hyaline, ca. 0.3 mm broad, entire, glabrous; 
receptacle concave, without resin; androphore pentagonoid, 2 mm long, 
L5-1.7 mm tall, L3 mm wide; stamens 18, L7-L8 mm long, the fila- 
ments flat, 0,3-0.4 mmm long, the anthers muticous, 1.4-1.5 mm long, 
0.5 mm wide, the apex emarginate, base cuneare, longitudinally dehiscent 
entire length; the connective medially darkened on both sides; pistillode 
absent. Pistillate flowers unknown. Fruit unknown. 

Type. VENEZUELA. Amazonas: Huachica, 1 1 km NE of San Carlos de Rfo Negro, 
1°58^N, 67^03^W, 120 m, 13 Nov 1977 (stam. fl), R. Uesner &J. Hall 3447 (holoiype: 
VEN; isoTYPES: MO, 2-shts), 

Paratypp^s: COLOMBIA. Caqui-ta: Araracuara, 0()^37'S, 72^24'W, 10 Dec 1991 (ster.), 
J. D/{!vemm)rden et aL 267 J (BRIT, CO AH, COL); Araracuara, Villa Aziil, Rfo Caqueta, 
left bank, in front of Sumaeta Island, 00^34'S, 72^08 W, 200-300 m, 4 Nov 1989 (srer), 
C. Umdmo et aL 1130 (DRFi; COAH, COL, JAUM, U). VENEZUELA. Ama/onas: 0.5-3 
km NE and E of San Carlos de Rio Negro, N of airstrip, 01^5 TN, 67^03^W, 120 m, 22 
Jan 1980 (ster.), II Lmmr 8576 (MO, NY, VEN). 

Distribution.— Kiu'^zon Basin of Venezuela and Colombia, in lowland 
areas of the Guayana Crystalline Shield, at 120-300 m elevation. 

Ecology and conservation status. — Ciusia grammadenioides occurs along 
steambanks on terraces just above floodline at forest margins on deep white 
sands. Because these areas are in such close proximity to major, frequently 
travelled river systems, this species should be considered threatened. 

Etymology. — The epithet "grammadenioides" refers to the resemble nee 
of the leaves of this species to those of several species of Cybianthus subge- 
nus Grammadema (Myrsinaceae). 

The decussate wings of the branchlecs apices and chartaceous leaves in- 
dicate a close relationship with the parapatric Ciusia dutdae. However, the 
broadly marginate petioles, short or obsolete peduncle, and sepals with 
scarious, hyaline and incised-erose margins clearly distinguish Ciusia 
grammadenioides from C\ duidae, 

3- Ciusia muitilineata Pipoly, sp. nov. (Fig. 2) 

Propter folia t^nuirer coriacea ad apiccm necnon basem atrcnuata, subter gland ulas 
punctacas puniccas induta atque lincas laticiferis atras praedita, non nunquam perianthii 
membra cucullata carinatast]iie ad C, dsymmvtrn-a Pipoly valde affinis sed ab ea foliis 
symmetricis necnon elliptkis (non asymmetricis necnon oblanceolatis), petiolis tenuis 1 — 
L5 (nee crassis 2-3 5) cm lon^is, denique cyma a eas ter (nee quater) ramificannbus facile 

Tree 3 m tall; branchlets subterete to weakly tetragonal, the corners 
rounded, not alate, 3-3^5 mm diam., subterete with age, glabrous. Leaves 
petiolate; blades chartaceous, elliptic, 8-13 cm long, (4. 0-) 5 -5. 5 (-6. 2) cm 
wide, apex and base acute, the base decurrenc on the petiole, symmetric, 


SiDA 16(4) 

Fig. 2. Clmin vuihsUmdtii Pipoly, A. Habit. B. Inflorescence. C. Staminatt flower buds. D. 
Petals and tUKlroerium. A-F, drawn from type. 

PiPOLY AND Graft, Clusia section Criuva in Guayana 657 

bearing round magenta secretory glands and numerous, conspicuous linear 
latex canals below, glabrous, midrib raised above and below, secondary veins 
numerous^ prominent above and below, united by a submarginal collecting 
vein, the margin entire, revolute; petioles marginate, ligulate, thin, 1 — 1.5 
cm long, glabrous. Staminate inflorescence terminal, erect, pyramidally 
paniculate, trichasial, 2.5—5 cm long, 3—7 cm wide, the flowers cymose; 
pedimcle 0.3— LO cm long; bracteoles 2, opposite, cartilaginous, ovate^ 1.5— 
1.6 mm long, 2—2.4 mm wide, apex acute to acuminate, cannate, w/o con- 
spicuous latex canalSj the margin scarious, entire, the border less than 0.1 
mm wide, glabrous; pedicels 2—4 mm long, tetragonal. Staminate flowers 
greenish -white; sepals 7, the lower 4 decussate, the upper 3 contorted, 
chartaceous, suborbicular, more or less equal in size, 2.2—2.4 mm long, 2— 
2.2 mm wide, apex obtuse, rugose medially toward the apex, slightly cari- 
nate, the margin scarious, entire, the border to 0.2 mm wide; petals 4, 
decussate, thickly carnose, without conspicuous latex canals, suborbicular, 
slightly carinate, the outer pair 4.0^.2 mm long, 3-2-3.4 mm wide, apex 
rounded, cucullate^ the margin entire, revolute, the inner pair 2,9—3/1 mm 
long, 2.0—2,2 mm wide, apex rounded, strongly cucullate, the margin en- 
tire, strongly revolute; receptacle concave, without resin; androphore 
rectanguloid, 0.8—1.1 mm high, 1.4—1.5 mm long, i.l-L2 mm wide; 
stamens 22-24, Ll-13 mm long, the filaments free, clavate, 0.3—0.4 mm 
long, broadly conic at base, the anthers muticous, obovoid, 0.8—0,9 mm 
long, 0.4—0.5 mm wide, apex emarginate, base obtuse, dehiscent by longi- 
tudinal slits for entire length; pisrillode absent. Pistillate flowers unknow^n. 
Fruit unknown. 

TvF>i:: VENEZUELA. AjMazonas, Depto. Ri'o Nc\^ro: Cerro de la Neblina, Camp IV, 15 
km NNE of Pico Phelps; 0^5 IN, 65^57'W, 7H() m; 15-lH M^ir 1984 (stam. fl), R. Lituier 
16649 (holotypc: VEN, lsotyphs: BRIT, MO, US). 

Distrihution. — Presumably endemic ro Cerro de la Neblina, Amazonas, 
Venezuela, 780 m, known only from the type. 

Ecology and comervatwn status. — C. ninltilineata is restricted to river can- 
yons of Cerro de la Neblina. Cerro dc la Neblina is very remote and lies 
within a national park and thus, is protected. 

Etymology. — The specific epithet refers to the numerous ^ conspicuous lin- 
ear latex canals of the leaf blades, which appear trans lucid upon drying, 

Clusia multilineata is closely related to C, asymmetrka Pipoly of Cerros 
Marahuaca and Aracamuni, but can mimediately be recognized by its el- 
liptic and symmetric leaves ^ thin petioles, and 3 -branched inflorescences. 

4* Clusia asyriinietrica Pipoly, sp. nov. (Fig. 3) 

Species notabilis haec en C, malt lit neat a di ni antequam refemntur ad ilia toliis 
asymmetricis necnon obianceolatis (non symmetric is nccnon ellipticis)^ petiolis crass is 


Sid A 16(1) 

Fic, 3- Chiiid ^symmetrica Pipoly. A. Habit. B. Staminate flower bud, C, Petals ant] 
antlrncciiim. A-R, drawn from type. 

2.0-3.5 (nee tenuis 1 .()~i ,5) mm longis, denique cyma a eas quater (non ter) ramificanribus 
starim separabilis. 

Tree 6 m call; branchlets tt^tragonLil, die comers nor alate, A-6 mm diam., 
remaining angular with age, glabrous. Leaves peciolace; blades thinly co- 
riaceous, obianceolate, (12-)1 5-20(-22) cm long, (4,8-)5-8(-9) cm wide, 
apex and base acute, the base decurrent on the petiole, asymmetric, bearing 
round magenta secretory glands and conspicuous linear latex canals below, 
glabrous, the margin entire; petioles trigonal, ligulate, chick, 2.0-3-5 cm 

PrpoLY AND GrafI", Clusia section Criuva in Guayana 659 

long, glabrous, S tarn in ate inflorescence terminal, pendent, pyramidally 
paniculate, 4-branched, 3—7 cm long, 7—10 cm wide, the flowers cymose; 
peduncle 03-1 cm long long; bracteoles 2, opposite, thickly cartilaginous, 
very^ broadly ovate, 3-6—3-8 mm long, 5-5.2 cm wide, apex rounded, me- 
dially rugose and prominently carinate, w/o conspicuous latex canals, the 
margin scarious, irregularly incised, the border ca. 0.4 mm wide, glabrous; 
pedicels 1 mm long, tetragonal. Staminate flowers white; sepals 4, 
chartaceous, orbicular, more or less equal in size, 4.0-4-2 mm long and 
wide, apex rounded, carinate and cucuUate, rugose medially coward the 
bii5Cj the margin scarious, irregularly incised, the border to 0.4 mm wide, 
glabrous; petals 4, imbricate, coriaceous, obovate, 4.8-5 mm long, 3-3^2 
mm wide, apex rounded, latex canals numerous, conspicuous, the margin 
entire^ flat; receptacle concave, without resin; androphore rectanguloid, 1.4- 
1.6 mm high, 3^8-4.1 mm long, 1.8-2 mm wide; stamens 18, 3-2-3.3 
mni long, the filaments basally connate forming a non-resinous mass of 
tissue 1.8-2.1 mm tall, apically free L4— 1,6 mm, subclavate, 0,^0.5 mm 
long, terete at base, the anthers muticous, ovoid, 0.8—1 mm long, 0.5-0.6 
mm wide, apex rounded to subemarginate, base acute, dehiscent by longi- 
tudinal slits for entire length, the connective not darkened; pistillode highly 
reduced, sparingly rcsiniferous, with hollow ovary and 5 styles and reduced 
stigmatic areas. Pistillate flowers and fruit unknowai. 

Type: VENEZUELA. Ama/onas. Depto. Atabapo: Cerro Marahuaca, "Sima" area, 
03^'43'N, 65^^30 W, 12()() m, 16 Oct 1988 (stam. fl), R. Lmner 24972 (holotypi- VEN; 
iS0TYPi;s: BRIT, MO). 

Pakaiypes: VENEZUELA. Ama/c)\as. Dcpro. RTo Negro: Cerro Aracamuni, ()1'^39^N, 
65^4{)'W, 250 m, 4 Nov I9B7 (stam. fl bud), il Lmner & G. CmiwvaU 22797 (MO, VEN), 
Deprc). Atabapu: affluent of (^afio Negro at junction with Ruj Cunucunuma, water source 
from Cerro Marahuaca, 1,000-1,200 m, 19 Oct 19fiR (stam. fl), H. Rfu/r/y^/fez 2873 (MY, 

Dhtrihution, — Endemic to southern Amazonas c^f Venezuela, on Cerro 
Marahuaca and Cerro Aracamuni, at 250-1,200 m elevation. 

Ecology and conservation status . — Clusia asyminetrka occurs in the ''si ma'* 
areas (sinkholes) of the upper plateaus of Cerro Marahuaca and the open 
"lajas" of Cerro Aracamuni. "Simas" have Inw-Ievel light regimes, high 
level moisture, but are shielded from the strong winds of the open areas on 
the tepui summits. The '"laja" formations described on the label of Lies ner 
& Cameualt 22797 are protected from winds by tall forests which surround 
them. While the environment on the tepuis is a fragile one, the southern 
tepuis are withm national parks and therefore, are pxroteccecL Despite the 
fact that this species is known from two vtry different elevational ranges, it 
is possible that exposure to w^ind, alternating periods of high humidity and 
aridity, concomitant with nutrient poverty are critical factors in determin- 
ing the species' success. 


Sum 16(4) 

Etymology. — The specific epithet refers to the asymmetric leaves, unique 
within the genus Cliisia. 

Within Cliisia section Criiwa {semii strkto), C. asymmetrka is easily dis- 
tinguished by its long, trigonal petioles and asymmetric leaves. The 4- 
branclied cymes wider than tall are also distinctive. 

5< Clusia melchiorii Gleason, Bull. Torrey Bot. Club. 58:403. 1931. Tvph. 

VENEZUELA. Amazonas: Cerro Duida, Desfiladero, 1,850 m, G. Tak^ 705 (holo- 

tvph: NY!; [Sotypi:: US!). 

CI mi a mvlch'ion Gleason, orth, var., ibid. 

Terrestrial, free-standing shrub or small tree to 4 m tall; branchlets 
subtercte, angulatc, (3— )7— 12 mm diam., not alate, glabrous. Leaves peti- 
olate; blades coriaceous or rarely thickly coriaceous, obvate to widely obo- 
vate, rarely oblong, (6.5-)8-10(-17) cm long, (2.8-)5.5-8(-ll) cm wide, 
apex broadly rounded to truncate , base cuneate, fully decurrent to petiole 
base, symmetric, without round magenta glands, linear latex canals incon- 
spicuous, nitid above, pallid below, midrib prominently raised above and 
below, secondary veins numerous, united by a submarginal collecting vein, 
the margin revolute, entire, glabrous; petiole broadly marginare, 3-10 mm 
long, glabrous. Inflorescence terminal, erect, pyramidally paniculate, (4.5-) 
8-ll(-19) cm long, (4.2-)6-l 1 cm wKk% the flowers cymose, in 7-9's; 
peduncle flattened, (2-)3.5-4.5 cm long, longitudinally ribbed, not alate, 
subtended bv 2 reduced leaves similar to vegetative leaves but 6-8 cm 
long, 2—3.') cm wide; secondary inflorescence bracts cartilaginous, oblate, 
2-5 mm long, 6-8 mm wide, apex broadly rounded, strongly carinate, the 
margin opaque, scarious, entire, glabrous; secondary peduncles similar to 
basal one, 5-10 mm long; bractcoles (4-)6-12(-l4), decussate, coriaceous, 
oblare, 2.5—3^5 mm long, 3.5^.5 mm wide, apex obtuse, the margin en- 
tire, hyaline, scarious, glabrous; pedicels tetragonal, 1-3 mm long. Stami- 
nate flowers yellowish-white; sepals 4, decussate, the outer coriaceous, ob- 
late, 3-5^ mm long, 4—4.5 mm wide, apex obtuse, carinate, the margin 
hyaline, scarious; petals ^6, carnose, obovate, 3-=3.5 mm long, 2.5-3 mm 
wide, apex broadly rounded, linear latex canals inconspicuous, the margin 
entire, opaque, not scarious, glabrous; androphore rectanguloid; stamens 
numerous, ca. 3-3^5 mm long, the filaments flat, 2.8-3 mm long, the an- 
thers muticous, oblong, ca. 0.5 mm long, 0.2 mm wide, apex rounded, 
base fused to filament, the connective not darkened. Pistillate flowers (bud) 
as in staminate, sepals identical; petals unknown; stami nodes 4, linear, 1 — 
1,5 mm long, the filaments 1-1,3 mm long, the antliers muticous, oblong, 
02—0.5 mm long, 0.1-0.2 mm wide, apex rounded, base fused to filament; 
pistil oblong to obovoid, 1,5—3 mm long, 2-^ mm wide; carpels 4; styles 
4, erect, connivent, 2-2.4 mm long; stigmas cuneiform, subpeltate, 0.6- 

PiPcH.y AND GrakE', Clusia section Criuva in Guayuna 661 

0.8 mm long and wide; ovules 4 per locule. Fruit ovoid, then fusiform at 
maturity, 10-13 mm long, 3-8 mm wide. 

DistrilmtiorL — Amazonian Colombia and Brazil, eastward to Surinam on 
sandstone mountains of the Guayana Crystalline Shield and its satellites, 
1,200-3,000 m. 

Ecology and conservation status, — Cliisia melcbiorji occurs in elfin forests, 
often dominated by Bonnetia sj^ecies, where it is a conspicuous element of 
the vegetation along margins of the forest on rockly slopes. Ic is a common 
species throughout its range and is not threatened at this time. 

Specimens examined. VENEZUELA. Ama/onas: Cano Verada, Campo M. Perez tu 
Camp Verada, 9()()-l,K)() m, 30 Jan 1951 (tr), /i M^igrnre el uL 31665 (NY, US, VEN); 
Cerrn Duida. Cerro Cuiebra, 1 ,5(K) m, 22-24 Apr 1949 (fr), II Ma^^mre & B. AUfi//hv. Jr 
29124 (NY, VEN), Culebra Valley, Rfn Cunuciinuma, 1,500 m, 19 Nov 1950 (fr). B. 
Mdy^/nre ei id, 295.35 (NY US, VEN); Cerro Guanay, ridye and savanna, W camp, 1,800 
m, 1 Feb 1951 (Ir), 6. t^UgHm ei <d. 31746 (NY VElN); Cerro Monche, Rio Ventuari, 
Ama/onas, 1,500 m, H Jan 1951 (fr), R Mai^/^irt ef a/. 30927 (NY VEN), Cerro dc la 
Neblina, Rfo Yuua, 2-^ km S (^f f^anip 3. 1,000 m, 24 Dec 1953 (fr), B. Mag/dre et aL 
3690^ (NY US, VEN), ciimbrc between Cumbre Camp and N escarpment, 1,700-1750 
m, 13 Jan 1954 (pisr. fl), R AUii/dre d al 3724S (NY, US, VEN), trail from camp to N 
escarpment, 1,800-1,900 m, 17 Nov 1957,(fr), B. Mag/dre er al 42124 (NY, US, VEN), 
upper basin of Canon Grande, above Sal to Grande, 1,900-2,000 m, 13 Dec 1957 (fr), B. 
MagNire et id, 42731 (NY US, VEN), E escarpment of Upper Cano Cirande Basin, 2,000 
m, 13 Dec 1957 (fr), B. Magmn d ai 42377 (NY, US, VEN), talus forest between Camps 
3 and 4, 700-1 ,300 m, 1 Jan 1958 (fr), /3. hUiguln et al. 42758 (NY, US,. YEN), headwa- 
ters of Cano Grande, SW jX)rtion, 1 ,900 m, 16-17 Oct 1970 (stam. fl), /. Steyejwark 103992 
(NY VEN); Planicie de /iiUicKiga, Rfo Tjtirico, 2,300 m, 10-15 Oct 1970 (stam. fl),/ 
S/eym/un-k 103847 (NY VEN); Neblina Massif, Camp 2, 2.8 km NE of Pico Phelps, 
0()"49'N, 65''59'W, 15 Apr 1984 (fr), B. Stem & A. Cemfy 1535 (BRIT, MO, NY EJS, 
VEN); Sierra Parima, 02^'27'2'i" N, 63''56'W, along Venezuelan-Brazilian border, 45 km 
NW of headwaters of Rfo Orinoco, 1,300 m, 18-23 May 1972 (stam. fi), / Steyermavk 
105958 (NY VI:N); Serranm Parii, Rfo Paru, Cano Asfsa, Rfo Ventuari, 6 km along W 
rim, 2,000 m, 4 Feb 1951 (stam. fl). It Cowan &J, Wimimk 31224 (NY, US, VEN), (stam, 
rt), II Onrav & J. Wimlack 3 1 257 A (NY US, VEN), (fr), R. Cmau & J. Wurdack 3 1 258 
(NY US, VEN); Cerro Si|^apo (Paniqiie), SE peak, W Mountain, 2,000 m, 20 Dec 1948 
(stam. fl), B. Magifire & L. P^ldi 277HI (NY, US, VEN), N escarpment, 1/1()0 m, 23 Dec 
1948 (stam. fl), B. Mag/^m & L. Poldi 27894 (NY US, VEN), Cano Grande, 1 km NW of 
Campo Grande, 1,100 m, 28 Dec !94cS (fr), B. Alagmre & L Polni 28014 (NY, US, VEN); 
Cano Profundo, l,iOO ni, 1 2 Jan 19l9 (stam. fl), B. Maguw & L Puliii 28302 (NY, US, 
VEN), 1 km above Intermediate Camp, (>00 m, 6 Feb 1949 (fr), B, Maginyi! & L. Poldi 
2879^ (NY US. VEN); Cerro Yapacana, Rfo Orinoco, summit, 1,200 m, 2 Jan 1951 (fr), 
B. Alag/m^e el aL 30642 (NY, US, VEN), broken cumbre at 1,200 m, 5 Jan 1951 (fr), B, 
Mag/nr^ d aL 30740 (NY US. VEN), summit, 03M5'N, 66^M5'W, 1.000-1,200 m, 5-7 
May 1970 (fr), y. Steyerm/rk 1 03 1 59 (NY VEN); Sierra Majgualida. NW sector, small 
valley along an up[>er tributary of Caiio Iguana, 05^30 N, 65''15'W, 2,000 m, 28 Feb-3 
Mar 1991 Or), P. Berry ei aL 4886 (BRIT, MO, VEN); P. Berr) ei aL 4926 (MO, VEN); 
Serranfa de Tapirapeco, Campo Taniacuari, stream crail from camp, 0UM4^N, 64^40'W, 
1,300 m, 10 Feb 1989 (fr), //■ Bed et aL 938 (BRIT, NY VEN); Cerro Yavf, 2,000-2,300 
m, 1-3 Mar 1947 (stam. fl), K P/jc//}s & C B. Hmbivck 78 (NY, VEN), Valley of Rio Corn- 


SiDA 16(4) 

CcH^t), W of SermiiKi dc YiLUije, plateau W of nver, 5'M2'N, 66°1()^W, 1,300 m, 7 Mar 
1987 (fr). /i. liolsi & R. Uesnir jj87 (MO, VEN); Sermnfa Yutaje, Rio Manipiare, NW 
ridge, 1,100 m, 1 1 Feb 1953 (fr), B. Maginn & C\ Maii^/mr .i'}l74 (NY, VHN). Bolivar: 
Distrito Piar, Aiiyan-rcpui', N portion of S section, W division of mountain, along K/o 
Chijrun, berwen N and S camp ar foot of "second wair\ K6()C) m, 11 May 1964 (fr), /, 
Steyermark 9.iH8l (NY, VEN), S -central region, heacKvaters of Rio Churun, 05 5 I 'N, 
62^^2'W, 200 m, 30 Mar 19H7 (stam. (1 biKl). B. Unls! i7H4 (MO. VEN), summit of 
central part of NW arn^, W division ol nioLMUain, along small creek S of the Jimmy Angel 
air|.lane, 1,800 ni, ^ May 1964 (fr),y, Stcyermark 95^38 {NY, VEN); Chimanta Massif, 
Central Section, NNW of Summit f^amp, 1,970 m, 19 Feb 1955 (fr), /. Shycrmirk & J. 
Win-iLh'k 94 J (NY, VEN), Torono-tepuf, N -hieing slopes on sun:imit above valley ol Ciino 
Mojado, 2,030-2, 1 50 m. 21 Feb 1955 (stam. il), /. Skytrmark &j. W/ndmk 1017 (NY, 
VEN), Vicinity ot Bluff Camp, at base ol W-facing bluffs of Torono-tepuf, 1,700 m, 2—4 
Jun 1953 (fr),/, Steyemuirk 75625 {NY VEN); Abacapa-tepuf, NW jiart of sumn^iit, 2,(K)()- 
2,125 ni, 14 Apr 1953 (fr)./ Steyerm/rk 73003 (NY, VEN), San^en-tepui, between Camp 
4 & 5, 1,500-1,750 ni, H Jan 1953 (stam. (i),J. WunUk 54092 (NY, VEN), Cerro 
Guaiquniima, Rfo Paragua, slopes behnv W escarpment, 1,600 m, 31 Dec 195 1 (fr), B. 
AidgHire 32891 (NY, VEN), summit, NE section, near cliifs, near headwaters of Rfo Cara|>o, 
05^-^59'N, 63"'25'W, 1,490- U50t) m, 25 May 1978 (fr),/ Skyermfrk ei .//. in320 (NY. 
VEN); Meseta dc Guait]uinima, S section, along Rfo Carapo, 8 km N oi Sal to (^ara|>o, 
05^^5rN, 63^33'W, 900 m, 6 Dec 1987 (fr), 0. Higher 12^37 (MYE, NY, VEN); Alto Rio 
Paragua, Rio Guaiia (Merevarf), near Brazilian ironrier, 20 Oct 1943 (pist. fl, fr), l\ Can/ofuf 
1036 (US, VEN); Meseta de Jaua, Cerro Jatia, summit of CW portion, fM5^' N, 64^^26'W, 
60 km NW of the Sanitary camp of the missKjn ol RTo Kanarakuni, 1,922-JJOO m, 21— 
27 Mar 1967 (stam. H), / Steyermark 98144 (NY, VEN), summit, {)4^4S'50" N, 64^34' 10 
W, SW part, along tributary of Rfo Marajano, 1,750-1,800 m, 22-^28 Eeb 1974 (stam. fi), 
J. Steyetv/ark e/ al /0936j (NY, VEN), E of camp, E of Rfo Marajano, 1,810-1, HHO m, 2S 
Eeb -5 Mar 1974 (stam. fl),/ Sfcytrmfrk et aL 109634 (NY, VEN); Summit t)t' SE-facing 
escarpment , E ol Cerro El Picaclio, N of Las Nievers and Las Chicharras, 15 km N of 
Tumeremo, vicinity of Debc^rah, Altiplanicie cfe Nuria, 000— 6'){) m, 5--8 Feb I96! (fr), /. 
Sttytrmark 89049 {NY, VEN), NW slopes of Ptan-tcpuf, 1,600-2,000 m, 17 Dec 1952 
(fr), B. AU/g///re&J. mm/dd 33903 (NY, VEN), ^4im)SS forest'^ cumbre. N valley, 1 JOO m, 
4 Jan 1952 (tr), /x Ah/i^/fire 32990 (NY, VHNj, along base of S-facing blufls, 2,410 m, 6 
Nov 1944 (pist. ill J. Skyermark 39903 (E. NY, US, VEN); Ilu-tepui, vicinity Camp No. 
1, K500 ni, 13 ^En- 1952 (fi9, «- AUr^nm 33391 (NY, VEN); [Jaipan-tquif, W peak, 
1,930 m, 4 Mar 1967 (stam. fl), T Koy./f^^.i & G. Af^osfhn 7482 (NY, VEN); Sierra Pakaraima, 
headwaters of Rin Paragua, along Venezuelan-Brazilian border. No. 15, 03''40'N, 63"00'W, 
1,400 m, 4^5 May 1973 (stam. ft), J. Steyermavk 107282 (NY, VEN); Cerro Rt>ramia, SW- 
facing slopes betw^een Rondt3n C^an^p and base of bluffs, 2J)40-2,22') m, 30 Sep 1944 
(stam. fl),/ Snyennark 38943 (E, NY, US, VEN), SW-facing slopes, Jan 1939 (fr), A. 
PiukMS 161 (NY, VEN). CjUY^ANA. Mazaruni-Potaro Region: Upper Mazarum R]\er 
Basin, Pakaraima Mts., along Karowrieng River, unnamed peak NW of Maipuri Ealls, 
05"43'N, 6(r08'W, 1 ,385 m, I3 Jun 1986 (fr), y. P//mly & K. Aifml 77 j / (BRG. NY, US), 
Headwaters of Karowrieng River, 06"12'N, 60'WW, 712 m, 15 Jun 1986 (bud), /. P//Wj 
& K. Alfred 77H9 (BRG, NY, US); N of Imhaimadai airstrip, 05^^43^N, 60°18'W, 51 1 m, 
15 Jun 1986 (fr),y. P//W) & K. A/Jm/ 782! (BRG, NY, US), Cuyuni-Mazaruni Region: 
Chi-Chi Mts., 4 km W of Chi-Chi Tails, on tributary leading N from Mazaruini River, 
()5"34^N, 6(rM5'W, 4^0 m, 18 Feb 1987 (tr),/ F^/m/y et al 10293. 10330 (EIX;, NY, 
US), upper Potaro-SiparLmi Region: summit of Mr Wokomung, 05^05'N, 59*^'50'W, 
1,650 m, 7 Jul 1989, (pise, fi bud), tl Buwn & G\ SiUNuels 9120 (l^RG, lUilX NY); 1,5 30 

PiPOiY AND Graff, Clusia section Criuva in Guayana 663 

m, 13 Jul 19.S9 (stam. fl bud), B. Boom & G. Samiids 9201 (BRG, BRIT, NY); iMt. 
Wokumun^, central summit plateau, 0.5-1 km S, 1—2 km E along cteek area and ridge, 
()5°()4'N, 59"52'W, 1,500-1 ,53{) m, 22 Feb iy93 (ft), T. Hmikel et al. 1535 (BRG, US). 
Upper Takatu-Upper Essequibo Region: S. Pakaraimii Mts,, 5 km E of Tipuru Village, 
Ureisha Mt summit, 04°1 1'N, 59°3rW, 994 m, 4 Mar 1992 (fr), B. Hoffman et ai 1184 
(BRIT, BRG, US); NW Kanuku Mrs., summit of Nappi Mt., 12 km S of Nappi Village, 
03''1S'N, 59°33'W, 750-950 m, H Feb 1993 (fr), B. Hojfimn & II Foster 5567 (BRG, US); 
S Pakaraima Mts, Kopinang Falls and Savanna, 900 m, 29 Aug 1961 (stam. fl), B. Magnire 
et al 45986A (FDG, K, NY, US), 31 Aug 1961 (stam. fl), B. Alagum et al. 46()38a (FDG, 
K, NY, US), 3 Sep 1961 (stam. H),, B. Magum et al. 4608 1 A (FDG, K, NY, US); Kanuku 
Mts,, Nappi Mc. 03°19'N, 59''34'W, 960 m, 1 1 Nov 19H7 (fr), h\. Jamm-Jmvk et al. 875 
(NY, U, US), SURINAM. Wilhelmira Gcbcrgte, Juliantops, 1,200 m, 3 Aug 1963 (stam. 
fl),7. Silmlz 10303 (BBS, NY, US), 3 Aug 1963 (pist. fl),/ Sc/mIz 10296 (BBS, NY, US); 
Juliana Top, I 5 km N of Uicie River, 03"36'-03"4rN, 56^30'-56°34'W, 1,000-1,2.30 m, 
18 Aug 1963 (stam. fl), H. lywm et al 54864 (BBS, NY, U, US); Tafelberg, escarpment 2 
km S of East Ridge. 1 Sep 1944 (stam. (1), B. ALni//jre 24585a (BBS, NY, US), 1 Sep 1944 
(pist. H), B. Magujre 24585 (BBS, NY, US), between Savanna VIIl and SW escarpment, 3 
Sep 1944 (fr), B. Maguire 2464! (BBS, NY, U, US); Bakhuis Mts, along Kabalebo and 
Coppenhame Rivers, 800 m, 25 Feb 1965 (fr), P. Flomimtz & P. Maas 3040 (NY, U). 
FRENCH GUIANA. Saul, S of Mt. Galbao, 13 Dec 1976 (ft), J.J. dn Cnninlie 5401 
(CAY, NY, P); Mont Galbao, 14 Dec 1976 (fr), S. Morr et al. 8770 (CAY, NY, P); Mont 
Galbao, SE peak, 03°35'N, 53°16'W, 700 m, 13 Sep 1994 (stam. fl bud), B. Boom 10863 
(CAY, NY); Mont Galbao, E sector, ()3°36'N, 53°17'W, 650 m, 15 Jan 19«6 {fr),J.J. De 
Granville 876! , 8762 (CAY, NY, P). BRAZIL. Amazonas; Plato da Serra Araca, SE por- 
tion of Serra Norte, OO^'Sl'N, 63"22'W, 1,150-1,250 m, 15 Feb 1981 (fr), /. doAmaral.J. 
Pipuly et al. 1593 (BRIT, INPA, MG, NY), 24 Feb 1984 (fr), A. S. Tavares et al. 139 
(INPA, MG, NY), Plateau of northern massif uf Serra Araca, 01°51-57'N, 63°21-22'W, 
N part of N slope, 1 ,400 m, 21 Feb 1984 (fr), G. Prance. J. Pipoly et al 29209 (INPA, MG, 
NY), W slope of southern massif, 00°40'N, 63"18'W, 1,000 m, 21 Mar 19S4 (stam. 1!),/ 
Pipfdy & G. Saiiii/eh 6864 (INPA, MG, NY, US); 0-3 km N of km 21 1 of Perimetral Norte 
Hwy, Pico Rondon, ca. 0]°32'N, 62^4H'W, 3 Feb 1984 (fr),_/. Pipoly et al. 66l4 (INPA, 
MG, NY). Terr. Roraima: Serra Sebang, Vista General, 1,525 m, 16-18 Dec 1954 (fr), B. 
iWagu'm 40308 (NY). 

Because of the unique flattened inflorescence rachis and peduncle, nu- 
merous flowers and numerous bracteoles similar to the sepals, C. melchhrii 
is most closely related to C. cardonae, but readily separated from it by the 
broadly marginate petioles, obovate to very widely obovate leaf blades with 
cuneace bases fully decurrent to the petiole base, and fruit ovoid at first, then 
fusiform at maturity. 

6. Clusia cardonae Maguire, Mem. New York Bot. Gard. 9:483. 1957. 

Type. VENEZUELA. Bolivar: Uaipan-tepui, Rio Carom', 1,700 m, 1-15 Feb 1948 
(stam. Il), K. Phelps & CM. Hhchaxk 412 (holotype: NY!; isotype: VEN). 

Trees to 4 m tall; latex yellow, copious; branchlets 5—8 mm diam., acutely 
tetragonal, the corners sharp, upon drying appearing narrowly alate, the 
wings less than 0.5 mm wide. Leaves petiolate; blades stiffly coriaceous, 
oblong, elliptic, or rhombic, (5.5-)7-9(-ll) cm long, (4-)6-8(-9-5) cm 


Seda 16(4) 

wide, apex obtuse to broadly rounded, base obcuse, noc decurrent on the 
petiole, symmetric, nicid above, pallid or rarely nit id below, without ma- 
genta glands, midrib elevated above and beknv, the secondary veins nu- 
merous, conspicuous above and belcnVj united by a submarginal connecting 
vein, linear latex canals inconspicuous, numerous, the margin revolute, 
entire, glabrous; petioles canaliculate 15-25(-28) mm long, glabrous. 
Sraminate inflorescence termitial, erect, pyramidally paniculate, 7-9(-n) 
cm long, (5-)6-8 cm wide, the flowers cymose; peduncle obsolete; sub- 
tended by two reduced leaves similar to vegetative leaves but obovare spathu- 
late, 2,5-2.8 mm long, K2-2 mm wide; secondary inflorescence bracts 
carnose, widely ovate, 2-2.5 mm long, 2.3-2.5 mm wide, apex obtuse, 
medially carinate, the margin entire, opaque, not scanous, glabrous; 
bracteoles 4—8, decussate, coriaceous, ovate, similar in size and shape to the 
bracts; pedicels tetragonal, 2-3^5 mm long, glabrotis. Staminate flowers 
yellow; sepals 4, decussatate, carnose, oblate to suborbicular, 2-2.3 mm 
long, 2.2-2.4 mm wide, apex broadly rounded, subcucullate, the margin 
entire, opaque, not scarious, glabrous; petals 4-5, the outer pair opposite, 
the inner decussate or imbricate, coriaceous, suborbicular, 5-6 mm long 
and wide, apex broadly rounded, cucuUate, linear latex canals inconspicuous, 
the margin entire, opaque, nor scarious, glabrous; androphore jxntagonoid; 
stamens numerous, 2-3-5 mm long, the filaments flat, 1-2 mm long, the 
anthers ovate, (18-1 mm long, apex rounded, the connective slightly ex- 
ceeding the thecae, base cordate, the connective medially darkened; pistil lode 
absent. Pistillate inflorescence as in staminate but jx dice Is 2-3 mm long; 
sepals 2'-2.5 mm long, 2.4-2.7 mm wide; staminodes numerous, 1-1.5 
mm long, the hlanients 0,9- 1^2 mm long, broadly triangular at base, con- 
n J vent, the anthers ovoid to oblong, ca. 0.3-6 mm long, apex broadly 
rounded, base obtuse, fused to filament apex; pistil subglohose, 2.5—3 mm 
long and in diam.; styles obsolete; stigmas narrowly cuneiform, I-1.5(-2) 
mm long, 0.5-0.8 mm wide, subpeltate, convex. Fruit subglobt}se, at first, 
dien globose, yellowish-green, 5-9 mm long and in diam. 

Dntribiilion. — Endemic to tlie Guayana Crystalline Shield, Clima ainhmat 
is known from the state of Bolfvar, Venezuela and nearby Guyana, at 920- 
1,825 m elevation. 

Ecology and ams^rvation status. — CI a si a card mute is a rare species, occur- 
ring on summits of tepuis which experience very little disturbance. It is 
locally common, and therefore is nor likely to be threatened. 

Rfprt S( nr;icive sfxcimens cxiimintd: Vh;NHZl;ELA. Bolfvar: Dtto. Piar: f^himunra- 
Massif, Aprada-tc-pui; 1,400-1,500 m, Au^^ 1950 (stani. fl), L Bcrnann 93! (MER, NY. 
VEN); savanna at font of Aprada-tepuu 05'^"^23'N, 62''27'W, 1,200 m, 5 May 1 9H7 (stan-i, 
fl), (I Ihiher 12098 (MYF; NY, VEN); Camarcaibarai-tcpu^ SW-tiLcmg shniiklcr. OV'^rN, 

Pjpoly and Graif, Clusia section Criuva in Cniayana 665 

62"()1'W, 1,80(}-1,825 m 22-24 May 1986 (fr),/ Sieywmark et aL 131998 (MO, VEN); 
Cerro Amuray-tepuf, W division of Los HtTmanos range, summit, 05^55'N, 62^1 5'W, 
1,030 m, 27 May 1986 (fr),/ Steyermark 132175 (MO, VEN); Ilu-tepui; ndge E of Mesa 
Grande, 1,650 m, 9 Mar 1952 (pist. fi), B, Maguire 333 hS, (stam. fl), B. Maguire 33319 
(NY, VEN); mesa S of Terekeyuren-tepuf, 4() km NE of the Kamarata Mission, 05*^5 TN, 
62^03' W, 1J80 ni, 15 Jan 1986 (fr), 0, Hr^btr & S. Gorzida 11135 (MYE, NY, VEN); 25 
km N of Luepa. abng El Dorado^Luepa road, 05^55'N, 6r25'W, 1,350 m, 2 Sep 1986 
(scam, ii), O. liaber & L llenuif^ciez 1 1721 (MYF, VEN); Uaipan-tepLif, Rfo Caroni, 2,7()0 
m, 26 Nov 1946 (stam. fl), F. Cardona 2U64 (NY, VEN), summit of W peak, 1,980 m, 4 
Mar 1967 (stam. fl), G. Agostini & T. Koyami 7458 (NY, VEN), between the W and E 
peaks of Uaipari, 1,500 m, 4 Mar 1967 (stam. fl), T Koyama & G. Agostjnj 7473 (NY, 
VEN); Uei-tepuf, between SE slope and summit, between Luepa and Cerro Venamo, vicinty 
of km 125, S of El Dorado, 1,100-1,300 m, 7 Mar 1962 (stam. fl), /. Steyermdrk & L 
Amtegukta 22 (NY, VEN); vicinity of km 129-130, S of El Dorado, NE of Luepa, 800- 
1,200 m, 6-1 1 Mar 1962 (fr),^. Suyermirk & f., Arhteg/mta 90 (NY, VEN); Cerro Venamo, 
along Guyana border, 1,400-1,500 m, 1 Jan 1964 {fr),^. Steyerm^rk et ai 92502 (NY, 
VEN), NW slopes, between road to campamento 125 and forest above waterfall, 1,100- 
1,300 m, 14 Apr I960 (fr),/ Stey^emiark & S. Nikfrm 112 (NY, VEN); ridge of Fila de La 
Danta, between campamento 125 and km 127, between Luepa and Cerro Venamo, 1,200 
m, 15-17 Apr I960 (fr),^. Sfeyermark & S. Nilssm 204 (NY, VEN). GUYANA. Upper 
Mazaruni River Region: Karowtipu Mountain, between camp and peak on W side of 
mountain, 05M5'N, 6()^35'W. 920-1,180 m, 24 Apr 1987, B. Boom & D. Gopaul 7705 
(BRIT, BRG, NY). Cuyuni-Mazaruni Region: Pakaraima Mountains, Kurupung River, 
top of Kamarau Falls, O6^^06'N, 60°2rW, 350 m, 19 Jul 1992 (pist. fl, fr), B. Hnffman 
2086 (BRG, BRIT, NY, US); Pakaraima Mountains, NE plateau of Mt. Ayanganna, 
05^23'N, 59''58'W, 1,500 m, 30 Oct 1992 (|Hst, fl), B, H^gman 3104 (BRG, BRIT, US; 
Mt. Ayanganna, easternmost peak, 05^25^N, 59^57'W, 1,350-1,380 m, 1 1 Mar 1987 (fr), 
/ Pipoiy et nL 11102 (FDG, NY, US). 

The coriaceous leaf blades with prominent secondary veins, erect, pyra- 
midally paniculace inflorescence, flattened inflorescence rachis, and numer- 
ous, small bracteoles similar to tlie sepals serve to indicate that Clusia cardonae 
is most closely related to C. mdchiorti. However^ C cardonae is readily sepa- 
rated by its canaliculate petioles, oblong, elliptic or rhombic leaves not 
decurrent on the petiole, sessile stigmas and globose fruits. 

7. Clusia hexacarpa Gleason, BuIL Torrey Bot. Club 58:403- 193 L Type: 

VENEZUELA. Amazonas; Cerro Duida, streamside at Central Camp, 1,800 m (pi^sc. 
fl), G Tate 565 (hcjlotype: NY!). 

CljLSia hexacarpa var. ptanttpinana Steyerm., Field iana, Bor. 28:387. 1952. syn. nov. 
Type. VENEZUELA. Bolfvar: Ptari-tepuf, S-facing slopes, vicinity of "Cave Rock,'* 

1,800 m, 4 Nov 1944 (pist. fl),^. Steyermark 59866 (holotypi:: F!; isotyhes: NY!, 

Free-Standing shrub to small tree, to 8 m tall; branchlets tetragonal, 5- 
7 mm diann, the angles acute to obrusish when dry, glabrous. Leaves peci- 
olate; blades cartilaginous, elliptic, 4-7 cm long^ 2,5^(^.5) cm wide, 
broadly rounded at summit, obtuse at base, not decurrent on the petiole, 


SiDA 16(4) 

symmetricv subniticl above, pallid and without round magenta glands be- 
low, midrib flat to slightly impressed above, prominently raised beknv, the 
secondary veins very inconspicuous, numerous, united by a very obscure 
submarginal connecting vein, the margin cncirc\ scanous, opaque, glabrous; 
petioles broadly marghiate, 6-15 mm long, glabrous. Staminace inflores- 
cence a single, terminal, pendent flower; peduncle 8-15 mm long; sub- 
tended by two reduced leaves, the leaves coriaceous, obovate, L5-2,5 cm 
long, L5 cm wnde, apex obtuse to rounded, base obtuse; bracteoles 2, car- 
tilaginous, suborbicular, 3^5-4 mm long and wide, apex broadly rounded, 
carinate, the margin entire, opaque, not scarious, glabrous. Staminate flowers 
{immature bud) white, suffused with pink; sepals 4, decussate, stiffly co- 
riaceous, the outer pair suborbicular, 6-7.5 mm long and wide, apex broadly 
rounded, linear latex canals numerous, conspicuous, the margin entire, 
opaque, not scarious, glabrous, the inner pair 5-6.5 mm long, 4.5-5^5 mm 
wide, apex broadly rounded, Hnear latex canals moderate, conspicuous, the 
margin entire, opaque, not scarious, glabrous; petals 6, the outer two op- 
posite, cartilaginous, obovate to obovate-spathulate and somewhat clawed, 
6-9 mm long, 3.5-5 mm wide, apex broadly rounded, cucullate, linear 
latex canals sparse, inconspicuous, the margin opaque, entire, not scarious, 
the inner 4 imbricate, like rhe outer ones but progressively smaller, to 4.5- 
5 mm long, 2.5-3 mm wide; androphore pentagonal; stamens numerous, 
ca, 3.5—4.5 mm long, the filament fleshy, flattened, ca, 2 mm long, broadly 
triangular and connivent at bise, the anther linear, ca. k5-2 mm long, 0.3— 
0.5 mm wide, triangular in cross section, apically truncate , basal ly obtuse, 
fused with hlament; pistillode obsolete. Pistillate flowers as in staminate, 
but peduncle 15-20 mm long; bracteoles 1.6-2 mm long; sepals cartilagi- 
nous, oblate, 8-10 mm long, 10-12 mm wide, apex broadly rounded, the 
margin scarious; persistent; petals cartilaginous, obovate-spathulate to some- 
what clawed, apex broadly rounded, linear latex canals numerous, highly 
conspicuous, (12-)l4-18 mm long, 0.6-0.8 mm wide; persistent in fruit; 
staminodes 9, rectangular in outline, flattened, 2.8—3.5 mm long, 1-1.5 
mm wide, apex truncate, acute, with vestigial anthers; carpels 6(-8); styles 
l-'2 mm long; stigmas carnose, subpeltate, del tare, 3-3-5 mm long and 
wide in flower, 4,8-5 mm long and wide on mature fruit, ovules numer- 
ous, horizontah Fruit ovoid, (2.5-)3-3:5 cm long, 1.5-2.5 cm diam. 

D/j7r//;/^//(^j^i'. ^Endemic to southern tepuis of Amazonas and Bolfvar, 
Venezuela, at 1,4()0— 1 ,800 m elevation. 

Ecology and anmrvcitmn stains. — Clnua hexacarpa is common in Chisui scrub 
forests on outcrops and expose, steep slopes of tepuis, and as such, is not 
threatened. It is found with numerous Cyclanthaceae, at the junction of 
the scrub forests with seepage areas, such as the Bromeliaceae patches which 

PiPOLY AND Gkaif, Cliisia section Cnuva in Guayana 667 

form colonies in poorly drained areas over rocks. Ic is nor considered threat- 
ened or endangered. 

Specimens examined: VFNEZIJELA. Aniazonas: Cerro Aracaniuiii, SLimmit, Proa 
Camp, OP32'N, 65''49'W, l,U)0 m, 26 Occ 1987 (pisr, fl. fr), II Uesnm' & G\ Cmmiali 
224m (MO, VliN), 28 Oct 1987 (fr), II Liesmr & G. Carmvali 22^75 (MO, VEN); Cerro 
Huachamacari. RiV) Cuiuicunuma, Left fork, Guin Dc Dios, 1 ,H()0 m, 8 Dec 1950 (pisr. fl), 
B. Magnm 30028, 30033, 30179 (NY, VCN), (sram. fl), B. Aianuire 30029 (NY, VEN), 
Depto. Rio Negro; Cerro cle la Neblina, 6.5 km SSW of base camp, S extension of range, 
(X)M7'N, 66^1 rW, l,6t)0 m, 18 Apr 1984 (fr), B. Siein el ai. 1651 (BRIT, MO, VEN); 
Cerro de la Neblina, S-face of Pico I^helps Massif. ()()"48'N, 66^00' W, 1,550-1, 650 m, 13 
Apr 1984 (fr), A. Geniry & B. Steui 46596 (MO, YEN); escar]inienr above Camp IV, 1,650- 
U75() m, 30 Dec 1953 (fr). B. Maguiy^ el aL 36990 (NY, VEN), NW cumbre, 1,950 ni, 30 
Dec 1953 (pist. H, fr), Ma^^an et al. 37003 , 37022 (NY, VEN), vicinity Cumbre Camp, 
1,800 m, 10 Jan 1954 (stam.fl). B. Magmre et .// 37144 (NY VEN), NW Camp, 1,800 m, 
12 Jan 1954 (fr), B. Magnire et aL 37247 (NY, VEN), trail immediately below N escarp- 
mcrir, 1,850 m, 17 Nov 1957 (sram. fi), B. l\Utg/i ire 42132 (NY, VEN), escaqmient above 
Canon CTrande, E of Cumbre Camp, t,H()0 m, 22 Nov 1957 (pist. fl), B. Magiure et al 
42169 (NY, VEN), headwaters of Canon Grande, SE portion, 1,900 m, 16-17 Oct 1970 
(scanr fl), 7, Steyerwark 103973 (NY, VEN), (pist, ft) J. Steyermark 103975 (NY, VEN); 
Serran/a Paru, Rfo Paru, Cano Asfsa, SW escarpment, 1 ,850 m, 17 Dec 1950 (fr), B- Maguhr 
etaL 33283 (NY, VEN), summit, W rim, 2,000 m, 2 Feb 1951 (fr), II Onv^n &J\ Wmxiack 
31 !98 (NY, VEN); Serranm Yiiraje, Rfo Manipiare, NW ridge, 1,500 m, 23 Feb 1953 (fr), 
B.& C, ALrgmre 35393 (NY, VEN), 1 ,400 m, 1 Mar 1953 (fr). B. & C\ Magmre 35419 (NY, 
VEN), Coro^Coro Drainage, 1 ,500 m, 2 Mar 1953 (frj, B. & C, Mag/nre 3^)500 (NY, VVM). 
Bolfvar: Chimanta-Massif, Chimanta-tepur, C section, above summit camp, 1 ,940 m, 4 
Feb 1955 (stam. fl), /. Sieyermark & J. Wnrdack 412 (NY, VEN), bordering zanjon above 
summit camp, between Middle and Upper Falls of Rfo Tirica, 1,925-1,940 m, 4 Feb 1955 
(pjst. fl), /. Steyermark & J. Wmxiack 592, 593 (NY, VEN), summit, along Cano Mojado, 
l^y^S5-l,91() m, 23 Feb 1955 (fr),/ Sleymnark & J. Wnrdack 1107 (NY, VEN); NW 
slopes of Churf-tepuf (Muru^tepuQ, 25 Jan 1953 (pist. fl, fr),J. W^m/ack 34198 (NY, 
VEN); Uaipiu>tepui, 1,900 m, 1-15 Feb 1948 (fr), K Pheips & C B. Hlfckock 394 (NY, 

Cimu/ hexacarpci is closely related to C, radiata, but easily distinguished 
by the larger bractcoles, fewer sepals and petals, and most notably, the 
periantii persistent in fruit. The type of subsp. ptarhepuiaiia Steyermark is 
notable only for its rose nniarkings on the petals and purple stigmas. In 
other species oi~ Cimia I have seen in the field, stigmas are often more purple 
when exposed to the sun. 

Maguire had intended to describe another subspecies of C- hexacarpa, 
bearing the subspccific epithet ''oao(arpa\ and several specimens were dis- 
tributed bearing that epithet. Flowever, Maguire never published the name, 
and study of those specimens indicate that population differs from others 
only in having eight instead of six carpels. Many more specimens were 
available to me than were to Maguire, showing that floral mcrosity can 
change significantly even within populations. 


SiDA 16(4) 

8. Clusia radiata Ma^uirc & Phelps, Roc. S(x\ Vcnez. Cienc, Nat. 1 1 1952. 

Typii. VENEZUELA. AxiAy.oNAs: Cerro Guanay, SE escarpment, 1 ,8()() m, 4 Feb 

19Xi (pist. fl fr), II AL^-unr, D. P/W/^.i, C /)\ HiidKock & G\ Bmkwski 31738 (holo- 
iTPLi: NY!; iscnvpi^s: F!, VEN). 

Free -standing shrub to small tree to 5 m tall. Branchlets tetnigonal, 6- 

8 mm diam., the angles obtuse, not alate. Leaves petiolace; blades cartilagi- 
nous, widely obovate, widely elliptic to suborbicuar, 6-10 cm long, ")-7 
cm wide, apex and base widely rounded, base not decurrent on the petiole, 
symmetric, nitid above, pallid and wirhour magenta glands below, midrib 
shghtly elevated above, prominently elevated below, the secondary nerves 
barely distinguishable above, numerous, united by a submarginal collect- 
ing vein, linear latex canals inconspicuous, the margin entire, opaque, some- 
what scarious, glabrous; petioles broadly margmate, 6-10 mm long, gla- 
brous. Staminace inflorescence unknown. Pistillate inflorescence a sin^de 
(rarely 3) terminal, pendent flower(.s); pakincle annulare, 1.5-3 cm long, 
subtended by 2 small leaHike bracts 1.3-'5 cm long, 1.2-3.5 cm wide; 
bractcole.s 2, carnose, oblate, 12-14 mm long, 7-9 mni wide, apex broadly 
rounded, carinare, the margins entire, opaque, scarious. Pistillate flowers 
white; sepals 4, decussate, carnose, very widely {)vate, 12-M mm long, 7- 

9 mm wide, apex broadly rounded, linear latex canals inconspicuous, promi- 
nently carinate, the margin opaque, scarious, entire, glabrous; petals 8, 
cartilaginous, the outer and inner identical in shape, progressively smaller 
acropetally, the outer 4 decussate, the inner 4 imbricate, very widely obo- 
vate, 20-25 mm long, 16-20 mm wide, apex broadly rounded, linear latex 
canals few, moderately conspicuous, the margin entire, opaque, not scari- 
ous; staminodes numerous, 3.8-4 mm long, the filaments Hat, 3-3.5 mm 
long, the anther linear, 0.3-0.5 mm long, apex truncate, base not distin- 
gLushable from filament, the connective darkened; pistil globose; carpels 
7-H); styles minute, 0.(S-I.() mm long; stigmas carnose, cuneiform, 
subpeltate, .3-1 mm long and wide m Fruit. Fruit ovoid, 3.5-4 cm long, 
3.0—^4.0 cm diam. 

Distribulion. — Endemic to Cerro Guanay, Amazonas, Venezuela, at 1 ,cS0()- 
2,000 m elevation. 

Ecology and comervatmi status. — Clinia radiatct occurs on rocky otitcrops 
along steep slopes. Cerro Guanay is a remote mountain and although noth- 
ing is known of the population biology of the species, it is likely that it is 
not threatened. 

Reprfsennitive specimen exumirH-d: VENFZUELA. Amazonas: Ctrro Gunmiy, sum- 
mi r, 2,000 m, 4 l-eb 1953 (fr), S. iWitgiiirv ct al 31749 (NY). 

Clmia radiata is most closely related to C hexacarpa, but is easily distin- 
guished from it by the wider leaves, longer bracteoles, uniform and more 
numerous sepals, and more numerous petals. 

PiPOi.Y ANi) Graff, Clusia section Criuva in G nay ana 669 

9- Clusia niaguireana Pipoly, sp. nov. (Icig, 4) 

Quoad petioios late marginacos, florei* solirarios vel 3, fructuni nvoidcLim, himinas 
cartilaginous C. hexacarpae valcle cithnis, sed ah ea pcciolis {)bs(5ktis vcl 6 (non 6-13) mm 
longis, floribus 3~f> (nee 1-3), axillaribus (nee terminalibus), petiilis suborbicularibus (nee 
orbicLilanbus vel obovatis) denique staminodiis 4 (non 8—9) pnieclarc distinguitun 

Shrub to small tree to 4("U)) m tall; branchlets tetragonal, strongly 
ridged but not winged, (4.5— ) 5— 9 ninn diam., glabrous. Leaves sessile; blades 
cartilaginous; very widely obovate, or rarely oblong, (3-5— )5—7(— 10.2) cm 
long, (2,3— )3— 5(— 8) cm wide, apex rounded to truncate, base obtuse, not 
decurrent on the petiole, symmerric, nitid above, pallid and without ma- 
genta glands below, midrib raised above and below, secondar}^ veins 20—30 
pairs, not or barely visible, glabrous, w/o visible latex canals, the margin 
thick, opaque, revolute, entire, glabrous; petioles obsolete to 7 mm long, 
deeply canaliculate, ligulate, glabrous, Staminate inflorescence axillary, 
pendent, a twice-branched cyme, 2.5-3 cm long, the peduncle 5-7 mm 
long, the pedicels tetragonal, 5—7 mm long, glabrous; bracteoles 2, carti- 
laginous, suborbicular to oblate, 4—4,2 mm long, 4.5-4.7 mm wide, apex 
rounded, basally rugose, carinate, slightly cucuUate, the margin entire, thick, 
opaque, glabrous; sepals 2, decussate to bracteoles, thinly coriaceous, ob- 
late, 6.3—6,5 mm long, 1—1 .2 mm wide, apex rounded, cucuUate, the latex 
canals obscure, marginate, the margin chartaceous, ca, OJ mm wide, opaque, 
entire, glabrous; petals 4, decussate in pairs and to the sepals, thickly carnose, 
dimorphic, latex canals conspicuous the outer pair sublinguiculate, 5,2— 
5-9 mm long, the claw 0,7—1 mm long, 2.1 mm wide, the limb oblate, 
4.5^.9 mm long, 5.7-5.9 mm w^ide, apex broadly rounded, the margin 
thick, {)paque, entire, glabrous, the inner pair unguiculate, 4-^.3 mm long, 
the claw 1-1.1 mm long, l.y-lA mm wide, the limb oblate, 3-3.2 mm 
long, 35-3-7 mm wide, apex broadly rounded, the margin undifferenti- 
ated, opaque, entire, glabrous; receptacle convex, cubic, 1.(^1.8 mm high, 
L 1-1.2 cm long and wide, w^ithout resin; androphore cubic^ stamens 26, 
equal in size, free, 1-1 J mm long, the fikmnents fleshy, flattened, free, 
0.2—0.3 nim long, the anthers oblong, 0,9-1 mm long, ca. 0.5 mm wide, 
apex emarginate^ base deeply cordate, the connective darkened, 
glabrous, longitudinally dehiscent over entire length; pistillode absent. Pis- 
tillate inflorescence pendent, a reduced cyme, L5-2(-2.5) cm long, the 
peduncle 2-3(-5) mm. long, the pedicels tetragonal, 2-3 mm long, gla- 
brous; bracteoles 2, cartilaginous, suborbicular to oblate, 2,1—2.2 mm long, 
2,4-2.5 mm wide, apex obtuse, basally rugose, carinate, slightly cucuUate, 
the margin entire, thick, opaque, glabrous; sepals 2, decussate to bracteoles, 
thinly coriaceous, orbicular, 6,4-6,6 mm long, and wide, apex rounded, 
cucuUate, the latex canals obscure, marginate, the margin chartaceous, ca. 
0.7 mm Wide, opaque, entire or sparsely incised, glabrous; petals 4, dec us- 


SlDA 16(1) 

Ficp. 4. Clifs'hi nutpfirmna Pipoly. Hiibic. B. Fruit. ^\. Pistil lart^ cyme. D. Sraniinate pedicel, 
and separated sepal, showing scarious niart^in. ii. Petals and androecium. A-C, drawn I rum 
ry[>e; O-Ii, drawn from Sttyermark 93342. 

sarc ill pairs and to the sepals, thickly carnose, dimorphic, latex canals con- 
spicLioLis, the outer pair suborbicular, 4.5—4.9 nim lon^ and wide, apex 
broadly rounded, rhe margin thick, opaque, entire, glabrous, the inner pair 
suborbicular, 4.2^.3 mm k)ng and wide, apex broadly roLinLk^d, the mar- 

Ptpc^lv and Grajf, Clusia section Criuva in Guayana 671 

gin undifferentiated, opaque, entire, glabrous; staminodes 4, resembling 
stamens, 2-2.4 mm long, the filaments flattened, L4-L6 mm long, broadly 
expanded at base, the sterile anthers orbicular, ca. 0.7 mm long and wide; 
pistil 10— carpel late, strongly ribbed, subglobose, 6-6.3 mm long, 5.6-5.8 
mm diameter, styles obsolete, stigmas attached apically and basal ly, ciinei- 
form, 2-2/1 mm long, L4-L6 mm wide, persistent. Fruit subglobose, 
L7-2 cm long, 1.4-L6 cm wide at maturity, strongly ribbed. 

Typi:v VENEZUELA, Bouvar: Dticj. Piar, Macizu del Chimanra, sector N-ccEicral of 
Chimiinta-tcpui, eastern heudwaterscabeceras of Cano Chimanra, 5^1H'N, 62''()9'W, 2,000 
m, 26-29 Jan 1983 (pisc. fl, fr),^. Sieymnark 127 980 (iiolotypu: VEN; isotvpes: BRIT, F, 
MO-2 shts, US). 

Paratypes: VENEZUELA. Bol(\'Ar: Saddle between Terekeyuren and Murisipan-tepuf, 
1 ,650 m, 22 Mar. 1987 (fr), B. HdM 3469 (BRIT. MO, US, VEN); Dtto. Piar, central & W 
part of saddle between Camarcaibarai-tcpuf and Tereke-Yurcn-cepui, 1,800-1,900 m, 
05^52 N, 62^0 rW, 23 May 1986 (fr) II Uesmr et al 21006 (BRIT, MO, US, VEN); 
Auyan-repui, summit of central portion of NE arm (W ran^e), between "Drizzly C^imp" 
and ^^Rio Lomira Camp,^' UH()()-1,H50 m, 3 May \9M (pisr. fl, fr),^. A. Sh'yvrmark 93442 
(F, NY, US. VEN), same general area, woods beside small creek among savannas S of Jimmy 
An^el crash site, 1,800 m, 7 May 1964 (stam. tl),/ A, Steymmirk 93342 (NY, US, VEN); 
central-NW section of E arm, Auyan^tepui, ()5"^57'N, 62^'25^W, 1,950 m, 27 Aug 1983 
(stam. fl), 0. Hukrd al. ^096 (MYF, NY, VEN); Uei-tepuf, between SE slope and summit, 
between Luepa & Cerro Venanio, vie. of km. 12^, S of El Dorado, 1,1 00-1 ,300 m, 7 Mar- 
1962 (fr),/ A. Steyermark & L Amtegrfivtd 20 (F, US, VEN); Macizo del Chimanta, Dtto. 
Piar, central-southern section, wide valley between NE border of Torono-tepuf and central 
section of Chimanta-tepui, S drainage, 5"l6N. 62^()9^W, 2J00 m, 11-15 Feb. 1985 
(stam. bud),/ PJlmly et al 7261 (MYF, NY, MO, VEN), (pist. fl, fr), 0. WibeKj. Pij^oly d 
al 10171 (MYF, NY, VEN), fsram. fl), (K H/z/kt. J. Pipoly d al. 10219 (MYF, NY, VEN); 
central-NW section of Chimanra-cepuf, E headwaters of Cano Chimanta, 05°i8'N, 
62°()9^W, 2,0(K) m, 26-29 Jan iy<S3 (pist. fl. fr), 0. H/^kr &l Steyermark 6934 (MYF, NY, 
VEN), (pist. fl, fr),/ Skymikfvk et al. I279H0 (NY, VEN); Abacapa-tepuf, BmnMiu ft)resr, 
NW part of summit. 2,125-2,300 ni, 13 Apn 1953 (fr),/ A. Steyermark 74861 (F, US, 
VEN); Abacapa-tepui; above 1st line sandstone bluffs, 2,()()0-2125 m, 14 Aj^r. 1953 (fr), 
/ A. Sfcyermark 75002 (BRIT, F, MO, US, VEN); Ajiacara-tepui, elfin forest formation on 
plateau of SE-tacing upper shoukk^r of Apacara-tepuf, 2,00{) m, 19 June 1953 (tr),/ A. 
Steyermayk 757/7 (F, NY, US, VEN); Agi-^araman-tepLn; SE-facing forested slopes below 
escarpment, l,SHO-^l,955 m, 26 Feb. 1955,/ A. Steyermark &J^J- WimLuk 1 166 (pist. fl, 
fr), (F, NY, US, VHN). GUYANA. Potaro-Siparuni Region: Kaieteur National Park, N 
of Menzies Landing, ()5"^N. 59''29'W, 400 m, 26 Jan 19H7 (fr),/ Pipoly & C. Gharlunran 
10029 (BRIT, BBS, CAY, FDG, K, NY R U. US); Pakaraima Mts., Mt, Wokuniung, 
summit ridge of Kamiewah Pinnacle NE to S Pmnacle, "Little Ayanganna,^' 05^0 1'N, 
59^^52'W, 1,550-1,650 m, 17 Nov 1993 (pist. fl), T Heukd ei aL 4493 (BRCi, BRIT. US). 

Distri/jutiom-^-Endcmic to the eastern tepuis of the state of Bolivar, Ven- 
ezuela, and adjacent Gtiyana, at (4()()-) 1,100-2300 m elevation. 

Ecoiogy and comervalion sUitia. — Clusui mcigairuma is restricted to rocky 
outcrops, usLially on overhanging ledges, and is subject to high winds and 
driving rains. I observed small parches of this species on the Kaieceur Pla- 
teau, but most of them were not fertile. It appears that the species has a 


Sir>A )6M) 

considerable range, and because it occurs in a very hostile habitat, it is 
mosr hkely not threatened. 

Ely uio logy,- — It is with great [pleasure that I dedicate this species ro the 
hite Bassett Maguire, prodigious fieldworker, student of neotropical 
Clusiaceae and tlie floni of the Guayana Highland during his long career at 
the New York Botanical Ciardcn. He conducted many expeditions to the 
most remote 1 oca H ties of rlie Guayana Highland, where he collected excel- 
lent specimens despite the harsh fiekl conditions. 

Clmia magnmirna is most closely related to Clnsia hexacarpa, but may i^^e 
distinguished from it by the axillary inflorescence with greater number of 
flowers, the subsessile leaves, suborbicukir petals and fewer stami nodes. 

10, Clusia opaca Maguire, Bot. Mus. Leafl, 15:62. 1951. Tvn:: BRAZIL. 

Aniaiionas: Patli between headsviitcrs nf Ira-I^^arape and headwaters of Igarape Abiu. 
affluent of Rio llirairu, -1-6 Jul 1^48 (stam. (1), II IL SdmlJes & ii Lopez 10 192 

(\[i)[.<)TYV\'.: NY!; is<yrYPi:: GH). 

C//Lua mhtctii SteyLTnr, Fieldiana, Bot. 28:391. 1952. syn. nov. Tvpn. VFNFiZUliLA. 
Ama:^onas: C^ano San Miguel, above mouth of Irhana, Guainfa, 125 m, 26 Mar 1942 

(pist, fl, fr), LL Willums }4H9t< (holotypI;: F!; jnotypi:: USD^ 

Free-Standing shrub to tree 6 m tall; branclilets subterete, 3^ mm diam., 
sparsely longitudinally ribbed, glabrous. Leaves petiolate; thickly coria- 
ceous, oblong or elliptic, (4.5-)7-9 cm long, 2.5-3.5{-3) cm wide, apex 
and base broadly rounded, the base not decurrent on the petiole, symmet- 
ric, nitid abc]ve, pallid and without magenta glands below, midrib im- 
pressed above, prominently raised below, the secondary veins numerous, 
inconspicuous, united by a submarginal nerve, linear latex canals mcon- 
spicuous, the margin revolute, entire, glabrous; petiole canaliculate, (0.6-) 
1-1.5 cm long, glahnxus. Staminate inflorescence terminal, pendent, a com- 
pact compound cyme. 6-]8-fl()\vered, 1.5-2 cm long, and wide; subtended 
by a pair of leaflike bracts, 1.5^ mm long, 01-12 cm wide, apex broadly 
rounded, base obtuse, the margins revolute, entire; peduncle ra. 3 mm 
long, subterete, glabrous; bracts carnose, oblate, 1.8-2.2 mm long, 2-2.4 
mm wide, apex rounded, carinate, the margin entire, opaque, not scarious, 
glal^rous; bracteoles 4, decussate, oblate, carnose, 1.8-2.2 mm long, 2-2.5 
mm wide, apex obtuse, carinate, the margin entire, opaque, scarious^ gla- 
brous; pedicels angulate, 3-7 mm long, glabrous. Staminate flowers white; 
sepals 4-5{-6), the outer opposite, decussate to the bracts, membranaceous, 
orbicular, 3-5 mm long, and wide, apex broadly rounded, linear latex ca- 
nals lew, conspicuous, the margin entire, hyahne, scarious, glabrous, the 
inner ones imbricate, 3^8^.2 mm long and wide, apex broadly rounded, 
the margin entire, hyaline, scarious; petals 4-5, coriaceous, oblong, the 
outer ones opposite, the inner imbricate, all similar in shape, acropetally 
decreasing m size, 4-6 mm long, 3.8-4,5 mm wide, apex broadly rounded. 

PfPOLY AMD Grai'F, Clusiii section Criuva in Guayana 67 


cucullate, the linear latex canals obscure, margins entire, opaque, not scari- 
ous, glabrous; androphore penragonoid, concave; stamens numerous, 0,9— 
L5 mm long, the filaments flattened, connate basally, the distal ones 0.2- 
0.3 mm long, the interior 0.5—0^7 mm long, the anthers linear, oblongs 
0,7-1 mm long, 0.2-0.3 mm wide, apex rounded, base obtuse; pistillode 
obsolete, not resin iferous, Pistillate inflorescence as in staminate, but 
bracteoles 2—2.5 mm long, 2.3—2.5 mm wide. Pistillate flowers as in stami- 
nate, but sepals 4-6, 4-4.5 mm long, 4.5-3 mm wide, petals 4-5 mm 
long, 3.3—4 mm wide; staminodia numerous, 0,8—1.2 mm long; filaments 
flat, linear, anthers barely differentiated, ca. 0.2 mm long, apex rounded, 
base not differentiated; pistil subglobosc, 3—5 mm long and wide; carpels 
5; styles subobsolete; stigmas sessile, orbicular, ca. 1 mm diam., ovules 
numerous- Fruit globose, 1.2-1.5 cm long and in diam, 

DistvibiftwrL — Western Amazon Basin of Brazil, Venezuela and Colom- 
bia, at 80— 160 m elevation. 

Ecology and conservation stains, — ijiisia opaca is endemic to "Amazonian 
caatinga," campinas, and ''Bana" formations, all of which are lowland subxeric 
areas ot deep white sands, often near black water rivers, but not 
subject to inundation (Macedo & Prance 1978; Prance 1979; Prance & 
Schubarr 1978). These environments all share essentially the same nutri- 
ent cycling regime, and van,' only in terms of local species composition. The 
white sands are coarse in texture, extremely well -drained, and derived 
from eroded tepuis. Clmia opaca is locally common, and because it occurs 
near the major black water rivers of the Amazon Basin, which support 
relatively heavy river traffic, it should be considered threatened. 

Common names. — "Pai-nan-ge" (Brazil, Maku language); "copei," "upihi/ 
"baniha/' 'Vupi'' (Venezuela). 

SpfCimcMis examined: COLOMBIA. Caqueta: Araracuara, santlsronc j^lateLiu behind 
military camp, ()()"i7% 72^24'^X/, 1 S Oct 1990 iii\J. D/avemmrdai & A. Ckef3l4 (BRIT 
COL, U). VENEZUELA. Depto. Atabapo: SL bank of the mitkllc part of Caiio Yagua at 
CucLinral cle Yi^iia, ().V\^6^N, r)6^^<i'W, 120 m, S M^iy 1979 (fr), G". Davuht^ tt id. 1736! 
(MO, MYF, NY, VEN); El Almidon, limit ()f Depts. of Atabapo and Casiquiare, Rfo Atacavi, 
slope 2, 03 04'N, 67=06'W, SO m, Nov I 9H9 ifr),J. Wazaj 869 (MUX PORT VEN); 
Near San Anrnnio, Alto Orinoco, alon^^ Rfo Orinoco. 1 20 ni, 15 Aug 1 9H2 (pisr. fl), f. 
iUfz el al. 3964 (MY, VLNf); R/o Guainfa, 14 Apr 1953 (pist. H), B, Alag//n-e &J. WimLck 
35642 (E MO, NY, US. VEN); savanna 5 km E of Maroa, 130 m, 6 Occ 1957 fstam. fl), II 
A^K/mt et di 41705 (F, NY, US, VEN), (pist. H biicl), II Ah^i^u/re et ai 41 706 (F, NY US, 
VEN) Maroa-Yavita road, between Rfo Gtiainfa and Cano Pimichrn, ca, 2 km beyond 
Maroa airport. 02 43'N, 67°38^W, H Oct 1978 (pist. fl, (r). H. Clark 6S63 (MO, NY, US, 
VEN), Maroa, Rfo Guainia, 127 m, 1942 (fr), !l WWh/m 14254 (i; US); Pimichfn, 1 2« 
m, 2 Jul 1942 (fr). LI. WiUhnu.i 14 183 (F US): sa^'a]lna 0.5- 1.5 km N of Puerro Colombia, 
opposite Maroa, Colombia, 130 m, 12 Oct 1957 (fr), II Alaj^//ire d uL 41843 (l\ NY, 
VEN); Depro, Casiqinarc. Rfo Gasiquiare, 10 km beyond the mourli and 5 km NE of 
camp, 28 Jan 1991 (fr), Al. Culclla i^t .//. 16H) (BRIT, NX VEN); Rfo Gasiquiarc, 162 kms 


Si DA 16(4) 

from tlu^ mouth, 3 Feb 1991 (ir), Al. Coklb ,t J. 1752 (BRIT. NY, VHN.)tmixed collec- 
tion wjth C ^it/fcluha/idn CJit)isy ex PI. ik TrJ, Cafiu San Miguel, sector "Las Tinajas, ' 
()2^l9^N, 66^4')' W, 160 m, 25 Apr 1991 (fr), G, Ay mem! 9 2 '57 (BRIT, PORT, VEN). 

CliLsici npaca is most closely related to C\ g/uryanae, burr easily disrin- 
^Liislu^d by the obtuse to broadly rounded leaf bases, the shorter peduncle, 
SLiborbicular sepals, coriaceous petsls, numerous stami nodes and sessile^ 
orbicular stigmas. 

IL Clusia guayanae Pipoly, sp. nov. (lig. 1) 

Species haec quoad periolns canal icularos, Hcres 6-18 in cyma Insidens, fnic rus globosiim, 
laevem, non costaroque, C npadw valdr arcre afhnis, sed ab ea laminis ad bascm obtusis vel 
hire roruiidacis (non acuris vel obuisis). pedunculis 5-12 (non 3-3) mm lon^^MS, sepalis 2 
(lion 4-6), nblacis (nee suborbicLiiaribus), petal is cart i lag inosis (ntjn coriaceis), scammodiis 
4 (non nim^erosis), denique srigmatibiis penragonis (non orbicularibus) perspicLie recedit. 

Shrub to 2 m; branch lets tetragonal, 3.5—5 mm diam., glabrous; latex 
white. Leaves peciolace; blades coriaceous, t)blanceolate to obovace or rarely 
suborbicular, (3-)4-6(--7) cm long, (1 ,5-)2-3,5(-4) cm wide, obtuse to 
truncate, base obtuse to rounded, nitid above, pallid and without magenta 
glands below; glabrous, midrib slightly raised above, prominently raised 
below, secondary veins numerous, at a steep angle from midrib ro a large 
submarginal collecting vein, ca. 0.3 mm h"om margin; petiole canaliculate, 
3-5 mm long, with a deep channel at base. Stami nate inflorescence termi- 
nal, a pendent panicle, 2-3 cm long, 2--3 cm wide, the branches cymose, in 
sets of 3; peduncle sc|uarrose 5-8(-12) mm long; secondary branch bracts 
2, carnose, depressed ovate, L8-2 mm long, 2.1-3 mm wide, apex broadly 
rounded, glabrous, margins entire, opaque; bracteoles 4, decussate, carnose, 
depressed ovate, 2,2-2. 1 mm long, 2.(S-2.S mm wide, aj^ex broadly rounded, 
glabrous, margins entire, opaque; sepals 6, coriaceous, decussate, the outer 
2, depressed-ovate cucullace, carinate, 4—4.5 mm long, 5-5.5 mm wide, 
a[^ex broadly rounded, [i:iargins entire, opaque, glabrous, the inner ones 
oblong, cucullate, 5-5.3 mm long, 4^.5 mm wide, apex truncate to widely 
rounded, the margin scarious, entire; petals 1, cartilagmous, oblong to 
widely oblong, cucullate, apex obtuse to truncate, 3*5-5 mm long, 2-3 
mm wide, margin irregular, o]")aque, rliick; androph(5re concave^ cuadrate, 
0,8-1 mm long, 1.4- L6 mm diam.; stamens 8, oblong, muticous, fila- 
ments and anthers undifferentiated, 0.8-1 mm long, 0.3-0^4 mm wide, 
apex rounded; pistil lode absent. Pistillate inflorescence as in staminate, 
but 1.5—2 cm long, 1.5 cm wide, secondary branch bracts 2, as in stami- 
nate; bracteoles 4, decussate, as m staminate flowers; sepals 6i, decussate, 
depressed ovate, the outer ones sepals 3.8—1 mm long, 4.2^.5 mm wide, 
inner ones 4.8-5 mm long, 4—4.2 mm wide, coriaceous, cucullate, cari- 
nate, apex broadly rounded, the margin scarious; petals deep pink, 4-5, 

PiPOLY AND GRAir, Clusia section Criuva in Guayana 


Fig, 5. CInsuj gimyanae Pipoly. A. Habir. B. Immature inflorescence. C. Androphore and 
andfoecmm, D. Bracts and sepals, H, Corolla. R Staminodia and ovary. G. Fruit, A-C, 
drawn from Steyermark & Hf^Lst 20^H4\ D-G, t^irawn from Huher 9}jiX 

carcilaginous, oblong, 5-7 mm long, 2-3 mm wide, cucullate, apex broadly 
rounded; staminodes 4, 1.8-2.2 mm long, thin, strap-likc% bases united by 
a flat tube 0.2 mm lon^, the filaments 1-L2 mm long, broadly ovate, the 


Si[^A 16(4) 

anchers widely oblong, 0.8-1 mm long, 0.6-0,8 mm wide, apex mucicous, 
with narrow^ longitudinal slits; pistil 5-carpellare, subglobostr, 2.7—3 nini 
long and diam., the stigmas peltate, subsessile, pentagonal. Fruit depressed- 
globose, 0.8-1 cm long, l-[.2 cm diam., pinkish yellow. 

Typi:. GUYANA. Cuvun[-Ma/.ar[ ni Rnc;um': Mr. Ayanganna, F side on scci]> slojies, 
5°27'N. 59^^^7'W, P25()-K3()() m. \2 Mar 1987 (pist. WlJ. Pifwly, G\ GbdrknTdu, G. 
Sammls.J. Chin 11 162 (noLcrr^Pi : FDG; isoivpks: BRIT, NY, US J. 

Pakajvpis. VFNKZUFLA. I^(jijvak: Dtto. Hltc's; Mllcj/o del Guaiquinima, central- 
NE section, slope drainin^y S; ^^'^-I'N; 63''42AV; \ 350 m. 1 Apr. 19X4 (pist. f1, fr). a 
ll//kr 9330 (MYF, NY, VEN); Drtu. Fiar, Central and western [xirt ol saddle between 
Camarcaibarai-tepui and lereke-Yuren-te|Hn, 05^52 N, 62^C)rW, 1,S()0-1,9()0 ni, 23 May 
19<S6 (stam. Il), R. Lmrm: er id. 20984 (MO, US, VEN); Clamarcaibarai-repuf, shoulder of 
W slope, easternmost tepnT of Aparanian-tepuf ran^^e, ()5'^'52'N. 62^'()rW, 1 ,S()() m, 2-1 
May 1986 (pisr. lb fr), B. Hobt et ai 2HH7 (MO, VEN); Auyan-tepur, summir, south- 
tenrral region, headwaters of R/o Clnirun. 05"^ PN, 62^32'W, 1,700 m, 29 Mar 1 9H7 (fr), 
II hfolst 373S (MO, VEN); plain in wesrernmt^st section of W arm of Aiiyan-tcpui^ 2^ km 
SE of Canaima, (,)r>'^()6'N, 62M3'W, [,()=)() m, 13 Nov 19^^■i (stam. i\), 0. Uuhcr 9728 

D'isfyihiftion. — Endemic to the easternmost tepuis of BohVar state, Ven- 
ezuela, and their satellites, at 1,2M)-I,90() m. 

Ecology dud coriservLftiori stat/LK^^-Cli/sid g/fayduae occurs in low, scrub 
cloudforest formations along edges of sandstone bluffs, dominated by Clmia 
melihiorii, C///s/d cm ss if olid Pbmch. & Triana, Bonnet id spp., numerous 
Myrtaceae and Rubiaceae. Its habitat has a hostile climate, with strong 
winds and rams. Because this S]'>ecies appears to inhabit the most inhospi- 
table of climates, it is not considered threatened. 

E/ywo/oi^y. — 'Hie sjuxific epithet, *guayanae" is in the Latin locative, for 
this member of the autochthonous flora of the Guavana Hit:hland. 

With canalicuate petioles, Howers in 6^-18-riowered cymes, and smooth, 
globose fruits without ribs, C giidydndc is most closely related ro C. updcd. 
However, C gaayande is easily separated from C. opdCd by the broadly acute 
to ohstuse leaf bases, longer peduncle, fewer, obbite sepals, cartilaginous 
petals, 4 stam i nodes and pentagonal stigmas on styles. 


Research for this treatment was great h' facilitated by the Missouri Bo- 
tanical Garden, w^hich provided loans and numerous gifts for determina- 
tion. Fieldwork was supjnjrted by the New York Botanical Garelen and 
Smithsonian Institutions duritig the senior authors tenure at tliose institu- 
tions. We thank Jon Kicketson, (Catherine Mayo and Roy Cummings (MO); 
June Cunningham and Lindsay Woodruff (BRIT) for technical assistance. 
Peter Stevens (A), Scott Mori (NY) and Barney Lipscomb (BRIT) jyrovided 
many useftil comments on the manuscript. Linda Ellis provided the excel- 

P[P(>LY AND GiiAir, Clusia section Criuva m CTuayana 


lent line illustrations in Figures 1, 4 and 5, while Juan Carlos Pinzon pro- 
vided Figures 2 and 3. R Mick Richardson (MO) faciUtated the particjpa- 
tion of the junior author in the project by arranging a one term study 
rotation in the senior author's laboratory 


Engier, a. 1888. Guttiftrae. In: C Martins, ecL Flora Brasiiiensis 1 2(1):3H2^74. 

1893. CImJa. In: Die Natiirlichen Pflanzenfamilien 3(6). Verla^ von Wilhelm 

Engelmann. Leipzig, 

Mac ][[>(), M and G. PKANfi-. 1978. Notes on the vegetation of Amazonia II. The dispersal 
of plants in An^a^onian white sand campinas: the campinas as functional islands. Brittonia 

MAGUfRH, B. 1979^ Guayana, region (if rlie Roraima Sandstone Formation. In: K, Liirsen and 
L. Holm- Nielsen, eds. Tropical Botany. Academic Press. New York. Pp. 223-238. 

PiPOLV, J. and A. Gram. 1995^ The genus C/usra sections Criuvopsis and Brachysremon 
(Clusiaceae) m northern South Aniirica. Sida 16:505-528. 

Pranch, G. 1979^ Notes on the vegetation of Amazonia III. The terminology of Amazo- 
nian forest types subject to inundation. Brittonia 31:26-38. 

Pranci-, G. and II. S(:i[i e^art, 1978, Notes on the vegetation of Amazonia I. A prelimi- 
nary note on the origin of the white sand campinas ol the lower Rio Negro. Brittonia 

Vi-sguii, J, 1893. Guttiferae, In: A. De Candolle, ed. Mongraj^hie Phanerogarum 8:1—669- 

numI'Ric;al llst of taxa 

1. Clusia duidac (ileason 

2. Clusia grammadenioides Pi poly 

3. Clusia midtilineata Pipoly 

4. Clusia asymmetrica Pi[>oly 

5. Clusia melchiorii Glcason 

6. Clusia cardonae Mai^uire 

7. Clusia licxacarpa Glcason 

8. Clusia radiata Maguire & Phelps 

9. Clusia maguireana Pipoly 
10. Clusia opaca Maguire 

1 I. Clusia guayanae Pipoly 

LIST iw h:xsif:cATAn 

The figures in parentheses refer to die numbeis from the numerical hst 
of taxa. 

Agostini, Cj & T Koyama 7458 (6). Amaral, h do; J. Pi}x)iy et ah 1593 (5). Aymard, G. 
9237 (10). 

Beck, H. et al. 938 (5). Bernardi, L. 931 (6) Berry, P et al, 4886 (5); 4926 (5). Boom, 
B. 10863 (5). Boom, B. & G. Samuels 9120 (5); 9201 (5X Boom, B. 6c D. Gopaul 7705 (6). 

Cardona, F 2064 (5); 1036 (5), Clark, PL 6863 (10). Collela, M, et al. 1752 (10); I6IO 
(10). Cowan, R. & J, Wurdack 31257A (5); 31258 (5); 31244 (5); 31198 (7). 

Davidse, G. et al. 17361 (10). Duivenvoordcn et al.. J. 2673 (2); 314 (10). 

Florschutz, R & R Maas 3040 (5). 

Gentry, A. ^ B. Stein 16596 (7). CiranviUe, J. J. De 5401 (5); 8762 (5); 8761 (5). 

Iknkel, T. Lt aL 4495 (9); 1535 (5). Hoffman, B. et ah 3104 (6); 3567 (5); 2086 (6); 
1181 (5). Hoist, B. 2887 (11); 3469 (9); 3738 (1 1); 3784 (5); 3387 (5). Huber, O. er a!. 
9728 (11); 9330 (11); I 1 135 (6); 8096 (9); 12137 (5); 12098 (6). Huber, O.J. Pipoly et 


Si DA 16(4) 

al. 10171 (y); 10219 (9). Huber, O, & 1,. Hernandez 1 1721 (6). Huber, O. <^j. Steyermark 

6934 (9). 

Irwin et al., H. 54864 (5). 

Jansen-Jacobs, M. 875 (5). 

Knyama, T & G. A^osrini 7473 (6); 74S2 (5). 

Liesner, R. et al. 8576 (2); 16649 (3); 17603 (1); 18533 (1); 209H4 (I 1); 21006 (9); 
24972 (4); 25135 d). Liesner, R, & G. Carnevaii 22489 (7); 22575 (7); 22797 (4). Liesner, 
R. & R. Hall 3447 (2). Londono, C. ec al. 1150 (2). 

Maguire, B. er al 24585 (5); 24585A (5); 24641 (5); 29124 (5); 30028 (7); 30033 (7); 
30179 (7); 30642 (5); 30740 (5); 29535 (5); 30017 (1); 30029 (7); 30927 (5); 3157S (8); 
31665 (5); 31746 (5); 31749 (8); 32990 (5); 33283 (7); 33318 (6); 33319 (6); 33391 (5); 
36905 (5); 36990 (7); 37003 (7); 37022 (7); 37144 (7); 37247 (7); 37248 (5); 41705 <10); 
41706 (10); 41843 (10); 42132 (7); 42169 (7); 42377 (5); 42731 (5); 42758 (5). Maguire, 
B. & C. Maguire 35174 (5); 35419 (7); 35500 (7); 35393 (7). Maguire, B. & L Poliri 
27781 (5); 27894 (5); 28014 (5); 28302 (5); 28797 (5); 32891 (5); 40308 (5); 459S6A (5); 
46038A (5); 46081 A (5). Maguire, B. ikj. Wurclack 33905 (5); 35642 (10). Mori, S. et al. 
8770 (5). 

Phelps, K. & C. B. Hitchcock 78 (5); 394 (7); 412 (6). Pinku.';, A. 161 (5). Pipoly, J. et 
al. 6614 (5); 7261 (9); 10293 (5); 10330 (5); 10330 (5); 1 1 102 (6). Pipoly, J. & K. Alfred 
771 1 (5); 7821 (5); 7789 (5). Pi|ioly, J. ik G. Gharbarraii 10029 (9). Pipoly, J. ^ C Samuels 
6864 (5). Prance, G., J. Pipoly et aL 29209 (5). 

Rodriguez, H. 2875 (4). Rui'z. T. et al. 3964 (10). 

Schultes, R. L. & F. Lopez 10192 (10). Schulz, J. 10303 (5); 10296 (5). Srein, R. & A. 
Gentry 1535 (5); 1651 (7). Steyermark, J. et al. 58945 (5); 59866 (7); 59903 (5); 74861 
(9); 75002 (9); 75005 (5); 75717 (9); 89049 (5); 9.3442 (9); 93542 (9); 92502 (6); 93538 
(5) 93881 (5); 98144 (5); 103992 (5); 103159 (5); 103847 (5); 10.5973 (7); 103975 (7); 
105958 (5); 107282 (5); 109363 (5); 109654 (5); 117320 (5); 127980 (9); 129648 (1) 
131998 (6); 132175 (6). Steyermark, J . >S; L. Anstcguicta 20 (9); 22 (6); 90 (6). Steyermark, 
J, & S. Nilsson 112 (6); 204 (6). Steyermark, J. & J, Wurdack 4l2 (7); 592 (7); 593 (7); 
942 (5); 1017(5); 1107 (7); 1166 (9). 

Tate, G. 429 (1); 705 (5). Tlwares, A. S. et al. 139 (5). 

Velazco, J. 869 (10). 

Williams, LI. l4l83 (10); 14254 (10); 11898 (10). Wurdack, J. 34198 (7); 34092 (5).